Diptera: Tipulomorpha: Limoniidae)

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Systematics Of The Neotropical Species Of Styringomyia Loew (Diptera: Tipulomorpha: Limoniidae)

Article in Zootaxa · August 2003 DOI: 10.11646/zootaxa.253.1.1

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Systematics of the Neotropical species of Styringomyia Loew (Diptera: Tipulomorpha: Limoniidae)

GUILHERME CUNHA RIBEIRO Pós-Graduação em Entomologia. Departamento de Biologia, FFCLRP-Universidade de São Paulo. Avenida Bandeirantes 3900, 14040-901. Ribeirão Preto, SP, Brazil ([email protected])

Contents

Abstract ...... 1 Introduction ...... 2 Materialandmethods...... 3 Morphology of the genus Styringomyia ...... 3 Key to the males of Neotropical Styringomyia species ...... 10 Styringomyia americana Alexander, 1914, nomen dubium ...... 11 Styringomyia dorsolineata Alexander, 1945, nomen dubium ...... 11 Styringomyia mystica Alexander,1945...... 13 Styringomyia youngi sp.n...... 16 Styringomyia atlantica sp.n...... 18 Styringomyia paulista Alexander,1946...... 19 Styringomyia simplex Alexander,1945...... 20 Styringomyia maya sp.n...... 22 Styringomyia amazonica sp.n...... 24 Styringomyia manauara sp.n...... 26 Phylogeny...... 27 Acknowledgements ...... 33 References ...... 33 Appendix ...... 35

Abstract

The Neotropical species of Styringomyia (Diptera: Limoniidae) are revised. S. americana Alex- ander, 1914 and S. dorsolineata Alexander, 1945 are considered nomina dubia. S. mystica Alex- ander, 1945, S. simplex Alexander, 1945 and S. paulista Alexander, 1946 are redescribed and illustrated in detail. Five new species, S. amazonica sp. n., S. atlantica sp. n., S. maya sp. n., S. manauara sp.n.andS. youngi sp. n. are described and illustrated. A cladistic analysis of the recog-

Accepted: 21 July 2003; published: 1 August 2003 1 ZOOTAXA nizable species (S. americana and S. dorsolineata excluded) is performed. A cladogram is produced 253 with the topology ((S. maya sp.n.(S. simplex + S. amazonica sp. n.)) (S. manauara sp.n.((S. mystica + S. youngi sp. n.) (S. paulista + S. atlantica sp. n.)))).

Key words: Styringomyia, Limoniidae, Diptera, taxonomy, systematics

Introduction

The limoniid genus Styringomyia was erected by Loew (1845) for S. venusta Loew, based on a single female specimen preserved in copal from an unknown locality of Africa. Soon after, Loew (1850) described S. gracilis from Baltic amber. It was only in the beginning of the twentieth century that the first recent Styringomyia, S. didyma Grimshaw, 1901 was described, from Hawaii. Since then, the knowledge about the world diversity of the genus has increased with the description of four additional fossil species (Cockerell, 1917; Cock- erell & Haines, 1921; Podenas & Poinar, 1999, 2001) and about 150 extant species from Neotropical, Afrotropical, Oriental and Australasian-Oceanian regions. Nevertheless, it is still Edwardss (1914) paper, dealing with no more than 23 species known in that time, the most comprehensive taxonomic revision made with the group to date. Nothing is known about the phylogenetic relationships among the species of the genus, and its affinities to other Limoniidae genera has not been well established. Edwards (1914: 206) called attention for the several peculiarities of structures that give to this genus a most isolated position among the Tipulidae (Tipulidae sensu lato, equaling Tipulomor- pha of the system used herein). Edwards (1924) remarked on a close proximity of Styrin- gomyia to the genus Teucholabis based on the morphology of the pupae without, however, giving further details. Alexander (1920) erected for Styringomyia the tribe Styringomyini, but later (Alexander, 1947), transferred it to the Styringomyaria, a subtribe of Eriopterini within the Tipulidae sensu lato. Hynes (1987, 1990), refused the necessity of placing the genus in any special tribe or subtribe, placing it in the already erected subtribe Eriopteraria Alexander in the Eriopterini. The only effort to elucidate the systematic position of Styrin- gomyia using phylogenetic methods was made by Oosterbroek & Theowald (1991), who placed the genus at the base of the so-called higher Eriopterinae clade, based on the presence, in the larvae, of reduced head capsule compared with the more massive head capsule of the lower Eriopterinae. Prior to the present paper, five species of Styringomyia have been described for the Neotropical region: S. americana Alexander, 1914 has been nominally recorded for Costa Rica, Colombia, Venezuela, Guyana, Suriname, Ecuador and Bolivia; S. dorsolineata Alexander, 1945 has been recorded for Ecuador; S. mystica Alexander, 1945, was recorded for Peru, and S. simplex Alexander, 1945, for Peru, Guyana and Bolivia. Finally, a single species, S. paulista Alexander, 1946, was recorded for Brazil, known only from its type locality in the southeastern region of the country (Alexander, 1947; Alexander & Alex- ander 1970).

2 © 2003 Magnolia Press RIBEIRO This paper presents a revision of the Neotropical species of the genus Styringomyia. ZOOTAXA Five new species are described, and the phylogenetic relationships among the species are 253 investigated using cladistic methodology.

Material and methods

The material of this study belongs to the collections of the Museu de Zoologia da Univer- sidade de São Paulo, São Paulo, Brazil (MZUSP), Instituto Nacional de Pesquisas da Amazônia, Amazonas, Brazil (INPA), National Museum of Natural History, Smithsonian Institution, Washington DC, USA (USNM), Carnegie Museum of Natural History, Pitts- burgh, PA, USA (CMNH), and The Academy of Natural Sciences, Philadelphia, PA, USA (ANSP). Informationonthespecimensexaminedaregiveninthesectionsdealingwiththe descriptions of their corresponding species. Data from specimen labels are in italics, and additional information and notes (e. g, geographical coordinates and depositaries) are given within parenthesis. Condition of preservation of holotypes is mentioned. Measurements were taken in microscopes with ocular reticules, and are given in milli- meters. Illustrations were made with drawing tubes attached to both stereoscopic and com- pound microscopes. Photographs were taken with a digital camera attached to a stereoscopic microscope. Descriptive terminology follows McAlpine (1981). Parsimony analysis was performed using PAUP (Phylogenetic Analysis Using Parsi- mony) version 4.0b10 (Swofford, 2002). Inapplicable, non-comparable, and unobserved character states were treated as missing data, and coded in the data matrix (Table III) as -, *,and?, respectively. Searching for the most parsimonious cladograms were made with the exact algorithm Branch and Bound. Characters were polarized a posteriori with root- ing following the out-group method revised by Nixon & Carpenter (1993). Character anal- yses were performed in MacClade version 3.08 (Maddison & Maddison, 1999). Details on decisions about character coding, ordering and weighting are given in the discussion of the results of the phylogenetic analysis.

Morphology of the genus Styringomyia

The diagnostic features of Styringomyia were redescribed by Edwards (1914, 1924). His descriptions, in several aspects reproduced by Alexander (1972), are accurate and relatively detailed, allowing the recognition of the main morphological attributes of the group. In the discussion below, the general characters of Styringomyia are dealt with, emphasiz- ing the morphology of the Neotropical species and structures not detailed described by Edwards (1914, 1924). The characters discussed hold constant through the studied taxa and will not be repeated in the individual species descriptions. The morphology of the male terminalia furnishes the most useful and decisive characters for recognition and com-

NEOTROPICAL STYRINGOMYIA © 2003 Magnolia Press 3 ZOOTAXA parison of species of the Neotropical fauna. The species descriptions are almost restricted 253 to the diagnostic characters of the male terminalia, and general metric data. Head and cervix. The head is somewhat dorsoventrally compressed, and a rostrum from 0.25 to 0.33 of total head length bear the mouthparts at its tip. The antenna (Fig. 1) is 16-segmented. The scape is stout and tubular in shape. The pedicel is ovoid, from 1.6 to 2.0 times shorter than the scape. The flagellomeres are ovoid and bear stout and relatively short spine-like hairs, which are little longer than each individual flagellomere. The length and width / length ratio of flagellomeres decreases gradually toward the tip of antenna. The maxillary palpus (Fig. 2) is 4-segmented, and the length of palpomeres increase from the base to the tip. The cervical sclerite has a peculiar shape: a basal piece, which is little longer than broad, articulates with a longer and narrow arm that articulates with the head at the margin of the occipital foramen (Fig. 3).

FIGURES 1-3. 1. Styringomyia mystica, holotype, antenna. 2. Styringomyia maya sp. n., male specimen from Costa Rica (Rivas, Talamanca), left palpus. 3. Styringomyia maya sp. n., male specimen from Ecuador (Manabi, Palmar), cervical sclerite. Specimens on slide. Abbreviations: ocp for, occipital foramen; ped, pedicel.

4 © 2003 Magnolia Press RIBEIRO Thorax. The thoracic anatomy of Styringomyia was only superficially described by ZOOTAXA Edwards (1914), and is described with much more detail here. The thorax (Figs. 4-6) is 253 dorsoventrally compressed. A medial longitudinal depression divides the pronotum dor- sally. The conspicuous pleural suture of the pronotum divides it laterally in anteponotum and postpronotum. The antepronotum is very enlarged, and has the anterodorsal margin protuberant. The postpronotum is less developed than the antepronotum. In the mesotho- rax, the episternal suture can be more or less conspicuous. The episternal cleft is very enlarged, and the anepisternum is restricted to a narrow area below it. Most of the mesepisternum is occupied by the katepisternum, and the great development of this sclerite makes the distance between the fore and middle coxae greater than that between middle and hind coxae. The mesothoracic pleural suture is usually strongly curved. The mesepisternum is about two times larger than the mesepimeron. The division of the mesepimeron is very conspicuous, being the anepimeron larger than the katepimeron. The sclerites of the metathorax are well delineated. The metaepisternum is clearly divided in metanepisternum and metakatepisternum. The metanepisternum is about 2.5 times longer than high, and encloses the posterior spiracle completely. The metakatepisternum is well developed and triangular in shape. The metaepimeron is narrow, inconspicuous, closely associated with the first abdominal segment. The laterotergite is not divided, and occupies an area similar to the area of the mesothoracic anepimeron. The legs are short and stout compared to other Limoniidae genera, and bear stout and spine-like hairs. The tarsus is 5-segmented. The basal tarsomere is much longer than the others, progressively shorter in length. The middle legs are shorter than the other two legs. Accurate proportions between the length of the fore, middle and hind legs are given by Edwards (1914).

FIGURE 4. Styringomyia dorsolineata, holotype, thorax, lateral view. Hairs not represented. Spec- imen on pin. Abbreviations: anepm, anepimeron; anepst, anepisternum; aprn, antepronotum; cx, coxa; kepm, katepimeron; kepst, katepisternum; ltg, laterotergite; mepm, metaepimeron; mtkepst, metakatepisternum; mtanepst, metanepisternum; mr, meron; mtg, mediotergite; pprn, postpronotum; sct, scutum; sctl, scutellum; tr, trochanter.

NEOTROPICAL STYRINGOMYIA © 2003 Magnolia Press 5 ZOOTAXA Wings. The wing venation of the Neotropical Styringomyia follows the general pattern 253 of the genus, described by Edwards (1914) and illustrated here in Figure 7. It is virtually useless for identifying species. Among the few characters showing variation, there is a lot of species polymorphism. One example is the shape of cell m1, that in conspecific specimens from the same locality, there may be a long petiole or lack this petiole at all. Almost all Neotropical species studied are polymorphic for this character. Pilosity and sexual dimorphism. Edwards (1914, 1924) called attention for the remarkable chaetotaxic pattern of the head and thorax in Styringomyia, which is unique within the Limoniidae. Although the general aspect of this chaetotaxic pattern is remarkably constant throughout the genus, differences in details of the arrangement and morphology of some bristles were found by Edwards (1924) to characterize groups of species from Afrotropi- cal, Oriental and Australasian regions. The Neotropical species fit well under the basic pattern described by Edwards (1914), and no significant variation was observed in the form and arrangement of bristles in different species. A representation of the thoracic chaetotaxic pattern in lateral and dorsal views are shown here in Figures 5 and 6.

FIGURES 5-6. Styringomyia paulista, holotype, thorax. 5. Dorsal view. 6. Lateral view. Specimen on pin.

FIGURE 7. Styringomyia paulista, holotype, wing. Specimen on pin.

The only sign of sexual dimorphism within the Neotropical species of Styringomyia can be observed in the pattern of the spine-like bristles of the fore femur and tibia. In males, these bristles are generally in greater number, longer, and more stout than in females. Female terminalia and male-female association. Thecercusandvalvearehairy, weakly sclerotized, and short compared with those of other Limoniidae genera. The cercus is triangular in shape, broad at base, and narrows gradually or abruptly toward tip. The profile of the ventral margin of the cercus is generally gently arched (as in Fig. 8), but is interrupted by a strong constriction near its base in S. simplex Alexander (Fig. 9).

FIGURES 8-9. Female terminalia of Neotropical Styringomyia species. Lateral view. 8. Allotype of Styringomyia paulista. 9. Paratype of Styringomyia simplex. Specimens on slides. Abbreviations: cerc, cercus; hyp vlv, hypogynial valve; tg 10, tenth tergite.

Since the characters that unequivocally allow species identification within the Neotro- pical fauna refer to the morphology of the male terminalia, in the absence of mating pairs among the studied specimens, male-female association can only be presumptive, and

NEOTROPICAL STYRINGOMYIA © 2003 Magnolia Press 7 ZOOTAXA based on co-occurrence. Several isolated female specimens deposited at USNM were asso- 253 ciated to one or another species by C. P. Alexander. However, the grounds on which such associations were made is not clear, and the possibility that some are erroneous cannot be discarded. Male terminalia. The male terminalia of Styringomyia has suffered a torsion of 180 degrees. The ninth abdominal sternite is dorsal, while the ninth and tenth tergites occupy a ventral position in the terminalia. In Edwards´ (1914) first account of the adult morphology of the genus, this aspect of the morphology of the terminalia was overlooked, and the author erroneously described the tergites as sternites and vice versa. A correct treatment of the dorsoventral orientation of the male terminalia, however, is given by Edwards (1924). The ninth sternite is generally triangular in shape, narrowing more or less abruptly from base to tip, whereas the ninth tergite is larger and more or less rounded. Articulating with the apex of the ninth tergite is the tenth tergite, which is weakly sclerotized and hairy. The tubular aedeagus is located ventrally in the interior of the aedeagal sheath, a large organ armed with large basal apodemes. The morphology of this sheath varies a lot within the genus as a whole, but there is little information in the literature about its morphology in different species. In the Neotropical and some non-Neotropical species, the aedeagal sheath has a dorsal crest of variable morphology, and two ventral branches of variable size and shape just below the opening of the aedeagus. Also, lateral lobes my be absent or present. The parameres are greatly developed and conspicuous in many Limoniidae genera, but cannot be recognized as separate structures in Styringomyia. It is possible that these structures have been lost in the genus, or became fused to constitute part of the aedeagal sheath. The morphology of the gonocoxite is also variable. The gonostylus is generally inserted at the apex of a dorsal extension of the basal part (sometimes, corresponding more or less with the basal half) of the gonocoxite. The posterior margin of this extension marks the point in which the gonostylus became narrowed, in some species, very abruptly. In most species of the genus, the gonostylus is divided into the inner and outer gonostyli. In all the Neotropical species, however, only one branch of the gonostylus is present, which is homologous with the inner gonostylus of other species. Different com- binations of extensions or protuberances are present in the gonostylus of the species of Sty- ringomyia. A study has been performed to determine the homology of these extensions among the Neotropical species, and their position, form and pilosity has proven to be informative for their interrelationships. In the Neotropical species, the number of extensions varies from one to four. For the set of species with three and four extensions (see the key and taxonomic part, below) the correspondence among them is not particularly difficult to assert (at least, in terms of hypothesis of primary homology sensu de Pinna, 1991), but this issue is more confusing for the species with a reduced number of extensions. For the species with four extensions, a reference number was given for each one, starting from the basalmost to the apicalmost (Figs. 10, 12 and 13, numbers I, II, III, and IV). For the species with less than four extensions (Figs. 11, 14-17), their form, relative positions and

8 © 2003 Magnolia Press RIBEIRO relations with other structures such as hairs, were used to determine their identity. The ZOOTAXA homology among the extensions of the gonostylus of the Neotropical species of Styringo- 253 myia is summarized in Figures 10-17.

FIGURES 10-13. Gonostylus of Neotropical Styringomyia species. Lateral and medial faces, respectively, on left and right sides of each figure. 10. S. mystica, holotype. 11. S. youngi sp. n., paratype. 12. S. atlantica sp. n., holotype. 13. S. paulista, holotype. The numbers refers to the homology system applied to the extensions of the gonostylus.

Coloration. The general color pattern is remarkably constant throughout the Neotropi- cal species of Styringomyia. Since the variation found in conspecific individuals are often barely distinguishable from that of different species, the study of coloration does not furnishes secure grounds for species identification. Furthermore, this general patter also occurs in species from other biogeographic regions, and may be plesiomorphic for the genus or, at least, plesiomorphic at the level of the Neotropical clade. The predominant color varies from yellowish to brownish. The scape, pedicel and palpi are darker brown to grayish. The lateral and posterior parts the thoracic tergum are darker than the medial parts. The thoracic pleuron and legs are more yellowish than the tergum. The fore femur has a brown band near its apex, and in some specimens, one or two lighter bands more proximad in position. In the meddle femur, a subterminal brown band may be absent or present. The hind femur is generally uniformly yellowish. The apex of tibiae is brownish. The last tarsomere is dark brown. The abdominal tergum is darker than the sternum, and has dark brown bands at the posterior parts of tergites. The wings are unspotted, except for the area around the vein r-m.

FIGURES 14-17. Gonostylus of Neotropical Styringomyia species. Lateral and medial faces, respectively, on left and right sides of each figure. 14. S. simplex, holotype. 15. S. maya sp. n., specimen from Ecuador (Manabi, Palmar). 16. S. amazonica sp. n., specimen from Ecuador (Francisco de Orellana). 17. S. manauara sp. n., paratype. The numbers refers to the homology system applied to the extensions of the gonostylus.

Key to the males of Neotropical Styringomyia species (based on the morphology of the gonostylus)

1. Stout spine-like hairs present (Figs. 10-13)...... 2 - Stout spine-like hairs absent (Figs. 14-17)...... 5 2. Gonostylus much longer than broad, bearing two long spine-like hairs; extension I greatly developed, basal in position (Figs. 10, 11)...... 3 - Gonostylus almost as long as broad, bearing at least three short spine-like hairs; extension I apical in position (Figs. 12, 13)...... 4 3. Four extensions present; apex of extension I broad (Fig. 10) ...... S. mystica Alexander - Only three extensions present; apex of extension I narrowed (Fig. 11)..S. youngi sp. n. 4. Extension II short, spine-like hair situated at its apex (Fig. 12)...... S. atlantica sp. n. - Extension II long, spine-like hair situated at its base (Fig. 13).....S. paulista Alexander 5. Two apical extensions present...... 6 - Only one bifid apical extension present (probably resulting from the fusion of extension III and IV); anterior margin of gonostylus almost straight; slender hairs in the lateral face concentrated at a protuberance near base of apical extension (Fig.

10 © 2003 Magnolia Press RIBEIRO 16)...... S. amazonica sp. n. ZOOTAXA - Three apical extensions present; extension III and IV closely jointed; gonostylus long 253 and slender (Fig. 17) ...... S. manauara sp. n. 6. Gonostylus more or less S-shaped; extension III relatively poorly developed; medial face with a long and relatively stout hair near base of extension IV (Fig. 14) ...... S. simplex Alexander - Gonostylus constricted at base; distal part of posterior margin expanded; extension III well developed (Fig. 15) ...... S. maya sp. n.

Styringomyia americana Alexander, 1914, nomen dubium

Styringomyia americana Alexander, 1914: 231 (description), Figure 1 (wing venation).

MATERIAL EXAMINED. Holotype. Sex unknown, Mallali, Demerara (ca. 6º46N, 58º10W), British Guyana, March 8, 1913, H. S. Parish (USNM). Most of the specimen pinned; left wing mounted on separate slide. DESCRIPTION. Dimensions: see Table I. REMARKS. Alexander (1914) superficially described this species based on two broken specimens lacking terminalia. Because the diagnostic features of the Neotropical Sty- ringomyia species refers to the morphology of the male terminalia, the taxonomic identity of the species-group taxon S. americana cannot be determined from its name-bearing type, and according to the International Code of Zoological Nomenclature (ICZN, 1999), it can be considered a nomen dubium. The holotype was identified by Alexander (1914: 231) as a male, although in the holotype label it is indicated that the specimen is a female. The information available does not allow the unambiguous determination of the sex of this specimen.

Styringomyia dorsolineata Alexander, 1945, nomen dubium

Styringomyia dorsolineata Alexander, 1945a: 419 (description).

MATERIAL EXAMINED. Holotype. Male, Ecuador, El Oro, Morro Morro (3º41S, 76º35W), 1500 m, 20-VII-1941, Laddey (USNM). Most of the specimen pinned together with allotype; right wing and some legs mounted on separate slide. DESCRIPTION. Dimensions. See table I. REMARKS. As for S. americana, the identity of this species cannot be identified from its name-bearing type due to the lack of information on the diagnostic structure of the male terminalia. Alexander (1945a) diagnosed this species by the relatively high degree of pigmentation of the first abdominal tergites, the general effect being a dorsal longitudinal

NEOTROPICAL STYRINGOMYIA © 2003 Magnolia Press 11 ZOOTAXA median stripe along the abdomen. However, the abdomen of the holotype is broken at the 253 level of the fourth segment, and its degree of pigmentation is not particularly different from the condition found, for example, in some specimens of S. maya sp. n., or in S. simplex, in which a longitudinal median stripe along the abdomen can be clearly observed. It was already stressed by Alexander (1945a: 419) in describing the species that: It is unfor- tunate that the hypopygium of the holotype male is lost, since the decisive specific character are to be found in this structure.

TABLE I. Selected measurements of Neotropical Styringomyia species (in mm)*. Maximum length Maximum width

Head Thorax Wing Gonocoxite Gonostylus Aedeagal Head Wing Gonocoxite Gonostylus sheath S. simplex

Holotype - 1.32 4.73 0.40 0.20 0.28 0.55 0.99 0.24 0.10

Range 0.70-0.72 1.32-1.60 4.68-4.73 0.39-0.41 0.17-0.20 0.28-0.39 0.40-0.55 0.99-1.10 0.21-0.24 0.09-0.10

Mean 0.71 1.47 4.70 0.40 0.19 0.33 0.46 1.00 0.22 0.10

n (2) (3) [2] [3] [3] [2] (3) [2] [3] [3]

S. amazonica sp. n.

Holotype 0.74 1.40 4.80 0.36 0.20 0.3 0.42 0.96 0.20 0.10

Range 0.66-0.74 1.40-1.42 4.80 0.36-0.38 0.20-0.22 0.28-0.30 0.40-0.42 0.90-0.96 0.20 0.10

Mean 0.69 1.41 4.80 0.37 0.21 0.29 0.41 0.94 0.20 0.10

n (2) (2) (3) [3] [3] [3] (2) (3) [3] [3]

S. mystica

Holotype 0.66 1.43 4.95 0.56 0.28 0.31 - 1.10 0.21 0.14

Range 0.60-0.66 1.40-1.43 4.60-4.95 0.50-0.56 0.28-0.34 0.28-0.31 0.32 0.86-1.10 0.16-0.21 0.10-0.14

Mean 0.62 1.41 4.78 0.53 0.30 0.30 0.32 0.96 0.19 0.12

n (3) (3) [1] + [2] [1] + [2] [1] + [2] [1] + [2] (2) [1] + [2] [1] + [2] [1] + [2]

S. paulista

Holotype 0.60 1.44 5.20 0.64 0.28 - - 1.10 0.16 0.20

Range ------

Mean ------

n (1) (1) (1) [1] [1] - - (1) [1] [1]

S. atlantica sp. n.

Holotype 0.88 - 4.31 0.56 0.21 0.24 0.66 0.92 0.23 0.19

Range - - 4.31-4.40 - 0.21-0.23 0.24-0.27 - 0.92 - 0.19-0.24

Mean - - 4.35 - 0.22 0.25 - 0.92 - 0.22

n [1] [1] [2] [1] [2] [2] [1] [2] [1] [2] ...... continued

Head Thorax Wing Gonocoxite Gonostylus Aedeagal Head Wing Gonocoxite Gonostylus sheath S. maya sp. n.

Holotype 0.64 1.46 5.86 0.52 0.20 0.32 0.30 1.18 0.20 0.08

Range 0.64-0.66 1.40-1.46 4.80-5.94 0.48-0.52 0.20 0.28-0.32 0.30-0.34 1.06-1.18 0.16-0.20 0.08

Mean 0.65 1.43 5.37 0.50 0.20 0.30 0.32 1.12 0.18 0.08

n (2) (2) (2) [2] [2] [2] (2) (2) [2] [2]

S. manauara sp. n

Holotype 0.52 1.12 4.20 0.54 0.20 0.26 0.30 0.88 0.18 0.06

Range 0.52-0.60 1.12-1.32 4.00-4.20 0.54 0.20-0.24 0.26-0.30 0.30-0.40 0.86-0.90 0.16-0.19 0.06

Mean 0.57 1.22 4.13 0.54 0.21 0.27 0.35 0.88 0.18 0.06

n (3) (3) (3) [3] [3] [3] (2) (3) [3] [3]

S. youngi sp. n

Holotype 0.62 1.40 4.80 0.50 0.22 0.32 - 1.06 0.20 0.08

Range 0.62-0.64 1.40-1.50 4.80-4.90 0.48-0.50 0.20-0.24 0.32-0.40 0.42-0.44 1.06-1.16 0.16-0.20 0.07-0.08

Mean 0.63 1.44 4.83 0.49 0.22 0.35 0.43 1.13 0.19 0.08

n (3) (3) (3) [3] [3] [3] (2) (3) [3] [3]

S. americana

Holotype 0.56 1.07 4.10 - - - 0.42 0.90 - -

Range ------

Mean ------

n (1) (1) [1] - - - (1) [1] - -

S. dorsolineata

Holotype - - 5.03 - - - - 1.18 - -

Range ------

Mean ------

n --[1]- - - -[1]- -

* The sex of the holotype of S. americana could not be identified unambiguously. For all other species, the measurements were taken from male specimens. The way of preservation of specimens may influence the dimensions of structures. The symbols ( ), [ ] and [ ] enclosing each n value iden- tify the specimens preserved on pin, slide, and glycerin, respectively. Since metric data are not used in this review to diagnose species, there was no concern in taking measurements from large series to give an exhaustive description of the range of the dimension of any of the measured structures.

Styringomyia mystica Alexander, 1945

Styringomyia mystica Alexander, 1945b: 241, Figure 7 (male terminalia); 261 (description).

FIGURES 18-25. Tenth tergite (male) of Neotropical Styringomyia species. 18. S. mystica, specimen from Peru (Madre de Dios). 19. S. youngi sp. n., holotype. 20. S. atlantica sp. n., paratype. 21. S. paulista, holotype (modified from Alexander, 1946). 22 S. simplex, specimen from Brazil (Mato Grosso, Diamantino). 23. S. maya sp. n., paratype. 24. S. amazonica sp. n., holotype. 25. S. manauara sp. n., paratype. Abbreviations: T9, ninth tergite; T10, tenth tergite.

MATERIAL EXAMINED. Holotype. Male, Peru, Junin, Satipo (10º19S, 74º23W), 800-900 m, February 5, 1940. P. Paprzychi (USNM). Part of the specimen pinned with allotype; other parts and terminalia mounted on slide. Allotype. Female,samedataof holotype (USNM). Paratype.1Male,Peru, Junin, Satipo, 800-900 m, February 5, 1940. P. Paprzychi (USNM). Additional material. 3 males and 5 females, PERU: Madre de Dios, Parque Manu, Pakitza, 11º56S, 71º18W, 250m, stream @ Tr1-13, IX-20-23-1989;

14 © 2003 Magnolia Press RIBEIRO MT, N. Adams (ANSP); 1 mating pair and 3 females, PERU: Madre de Dios, Parque ZOOTAXA Manu, Pakitza, 11º56S, 71º18W, 250m, stream @ Tr.2-12, IX-16-1989, LT, Gelhaus & 253 Epstein # 440 (ANSP); 1 female, PERU: Madre de Dios, Parque Manu, Pakitza, 11º56S, 71º18W, 250m, Qda. Claro @ Tr1-13, IX-8-1989, at dusk, J. Gelhaus # 435 (ANSP).

FIGURE 26. Styringomyia mystica, holotype. Aspect of part of terminalia. The arrow indicates the short and curved hair at the small protuberance near apex of ventral margin of gonocoxite. Speci- men on slide. Abbreviations: aed, aedeagus; aed sh, aedeagal sheath; goncx, gonocoxite; gonst, gonostylus; vb, ventral branches of aedeagal sheath.

DESCRIPTION. Dimensions. See Table I. Morphology. Ninth sternite: trilobulate; medial lobe smaller than lateral lobes, narrowing gradually toward tip. Tenth tergite (Fig. 18): posterior margin rounded. Gonocoxite (Figs. 26-27): distal half conical, abruptly narrowed, bearing two long and stout hairs at apex, and a stout, shorter and curved hair at an apical protuberance of ventral margin; other hairs slender, variable in length, the longest

NEOTROPICAL STYRINGOMYIA © 2003 Magnolia Press 15 ZOOTAXA along the lateral margin. Gonostylus (Figs 10, 26-27): with four extensions; extension I 253 (Fig. 10, I) very enlarged and sclerotized, bearing a long and a stout spine-like hair; extension II (Fig. 10, II) weakly sclerotized, shorter than the others, bearing long and stout spine-like hair at apex; extension III (Fig. 10, III) longer than the others, strongly sclerotized; extension IV (Fig. 10, IV) more slender than the others, strongly sclerotized. Aedeagal sheath (Figs. 26-27): a pair of lateral lobes present; dorsal crest three-pointed; medial point posterior in relation to the others; ventral branches long, narrowing gradually toward tip. Female cercus: narrowing abruptly toward tip; ventral margin gently curved, not constricted.

FIGURE 27. Styringomyia mystica, male specimen from Peru (Madre de Dios). Three-dimensional structure of terminalia, lateral view. The arrow indicates the short and curved hair at the small protuberance near apex of ventral margin of gonocoxite. Abbreviations: aed sh, aedeagal sheath; goncx, gonocoxite; gonst, gonostylus; S9, ninth sternite; T10, tenth tergite.

Styringomyia youngi sp. n.

MATERIAL EXAMINED. Holotype. Male, Ecuador: Zamora-Chinchipe, 84 km NW Zamora, mouth Rio Sabanilla (ca. 4º05S 79º01W), 1420m, 1 November 1987, C. Young, R. Davidson, J. Rawlins (CMNH). Pinned specimen, with dissected terminalia inside microvial with glycerin pinned with the specimen. Paratypes. 4 males and 3 females, same data as holotype (CMNH).

16 © 2003 Magnolia Press RIBEIRO ETYMOLOGY. This species is named after Dr. Chen Young (Carnegie Museum of ZOOTAXA Natural History, Pittsburgh, USA), one of the collectors of the type specimens, and the per- 253 son who made the Styringomyia specimens of CMNH collection available for this study. DESCRIPTION. Dimensions. See table I. Morphology. Ninth sternite (Fig. 28): narrowing gradually toward tip. Tenth tergite (Fig. 19): posterior margin rounded. Gonocoxite (Fig. 28): cylindrical proximally; distal part enlarged, bifurcated; dorsal branch with ca. five long and stout hairs along most distal parts of dorsal margin to apex; ventral branch with a ventral protuberance baring a usually short, curved and stout hair, and two long and stout hairs at apical margin; other hairs long and slender. Gonostylus (Figs. 11, 28): with three extensions; extension I weakly sclerotized, curved, narrowing gradually toward tip, bearing long and stout spine-like hair at its base; extension II (Fig. 11, II) weakly sclerotized, shorter than the others, bearing long and stout spine-like hair at its apex, extension III (Fig. 11, III) more sclerotized than the others extensions, curved, narrowing gradually toward tip. Aedeagal sheath (Fig. 28): a pair of large lateral lobes present; dorsal crest three-pointed; medial point bidentate, posterior in relation to the others; ventral branches long, narrowing gradually toward tip. Female cercus: narrowing abruptly toward tip; ventral margin gently curved, not constricted.

FIGURE 28. Styringomyia youngi sp. n., holotype. Three-dimensional structure of terminalia, dorsolateral view. The arrow indicates the short and curved hair at the small protuberance near apex of ventral margin of gonocoxite. Abbreviations: aed, aedeagus; aed sh, aedeagal sheath; goncx, gonocoxite; gonst, gonostylus; S9, ninth sternite, vb, ventral branches of aedeagal sheath.

NEOTROPICAL STYRINGOMYIA © 2003 Magnolia Press 17 ZOOTAXA Styringomyia atlantica sp. n. 253 MATERIAL EXAMINED. Holotype. Male, Brazil, Bahia, CEPEC-CEPLAC, Itabuna (14º48S, 39º18W), cacao leaf litter, 1970-72. J. A. Winder (USNM). Entire specimen mounted on slide. Paratype. Male, same data of holotype (USNM).

FIGURE 29. Styringomyia atlantica sp. n., holotype. Aspect of part of terminalia. Specimen on slide. Abbreviations: goncx, gonocoxite; gonst, gonostylus.

ETYMOLOGY. The name of this species refers to its habitat at the Brazilian Atlantic Rain Forest. DESCRIPTION. Dimensions. See Table I. Morphology. Ninth sternite: narrowing gradually toward tip. Tenth tergite (Fig. 20): posterior margin V-shaped. Gonocoxite (Fig. 29): conical, narrowing gradually toward tip; short and stout hairs along posterior part of medial margin to apex; other hairs slender, the longest along lateral margin, the shortest proximally on medial margin. Gonostylus (Figs. 12, 29): almost as long as broad, with four extensions; extension I (Fig 12, I) strongly sclerotized, bearing two apical spine-like hair; extension II (Fig. 12, II) shorter than the others, strongly sclerotized, bearing a short apical

18 © 2003 Magnolia Press RIBEIRO spine-like hair; extension III (Fig. 12, III) approximately triangular in shape; extension IV ZOOTAXA (Fig. 12, IV) rounded; slender hairs concentrated near base of extensions III and IV and 253 near the center of the medial face of gonostylus. Aedeagal sheath: with two lateral protuberances, the most dorsal rounded, the most ventral longer than broad, oblique in position; details of dorsal crest not visible in the available specimens; ventral branches short.

Styringomyia paulista Alexander, 1946

Styringomyia paulista Alexander, 1946: 7 (description); 9, Figure 8 (male terminalia).

MATERIAL EXAMINED. Holotype. Male, Brazil, São Paulo, Juquiá (24º48S, 47º36W), 300 m, February 22, 1941, Travassos (MZUSP). Part of specimen pinned; terminalia mountedinplasticslidepinnedwiththespecimen.Allotype. Female, same data of holotype (USNM).

FIGURE 30. Styringomyia paulista, holotype. Aspect of gonocoxite. Specimen on slide. Abbrevia- tion: goncx, gonocoxite.

DESCRIPTION. Dimensions. See Table I. Morphology. Ninth sternite: narrowing gradually toward tip. Tenth tergite (Fig 21): posterior margin slightly trilobulate. Gonocox- ite (Fig. 30): conical, narrowing gradually toward tip; relatively short and stout hairs along distal parts of medial margin to apex; other hairs slender, the longest along lateral margin,

NEOTROPICAL STYRINGOMYIA © 2003 Magnolia Press 19 ZOOTAXA the shortest proximally on medial margin. Gonostylus (Fig. 13): almost as long as broad, 253 with four extensions; extension I (Fig. 13, I) strongly sclerotized, bearing three apical spine-like hairs; extension II (Fig. 13, II) longer than the others, strongly sclerotized at apex, bearing a short subapical spine-like hair; extensions III and IV (Fig. 13, III and IV) approximately triangular in shape, similar in length; slender hairs concentrated near base of extensions III and IV and near the center of the medial face of gonostylus. Aedeagal sheath: with two lateral lobes, the most dorsal rounded, the most ventral longer than broad, oblique in position; details of dorsal crest ventral branches not visible in the available material. Female cercus (Fig 8): narrowing abruptly toward tip; ventral margin gently curved, not constricted.

Styringomyia simplex Alexander, 1945

Styringomyia simplex Alexander, 1945b: 241, Figure 9 (male terminalia); 263 (description).

FIGURE 31. Styringomyia simplex, holotype. Aspect of part of terminalia. Specimen on slide. Abbreviations: aed, aedeagus; aed sh, aedeagal sheath; goncx, gonocoxite; gonst, gonostylus; vb, ventral branches of aedeagal sheath.

20 © 2003 Magnolia Press RIBEIRO MATERIAL EXAMINED. Holotype. Male, British Guyana, Bartica (6º25N, 58º 37W), ZOOTAXA January 28, 1913, H. S. Parish (USNM). Part of specimen pinned; other parts and termi- 253 nalia mounted on slide. Paratypes. 1 male, Peru, Junin, Satipo, Jauja (10º19S, 74º23W), 800-900 m, XI-10-1940, Paprzychi (USNM); 1 male and 1 female, Peru,Junin,Satipo, Jauja, 800-900 m, IV-5-1941, Paprzycki (USNM). Additional material. 1 male, Peru, Huanuco, Fundo Sinchono, Cordillera Azul (8-10ºS, 74-76ºN), 1500 m, VIII-4-1947, J. M. Schunke (USNM); 1 male, Peru, Loreto, Cerra Azul, Rio Ucayali (8-10ºS, 74-76ºN),IV- 30-1947, J. M. Schunke (USNM); 1 male, PERU: Madre de Dios, Parque Manu, Pakitza, 11º56S 71º18W, 250m, stream @ Tr1-13, IX-20-23-1989; MT, N Adams (ANSP); 2 males and 1 female, Brazil: Mato Grosso, Diamantino (14º25S , 56º29W), Facienda São João, 5. II. 1981, 450m, C. Young (CMNH); 1 male and 1 female, Brazil: Mato Grosso, Diaman- tino (14º25S , 56º29W), Facienda São João, 6. II. 1981, 450m, C. Young (CMNH).

FIGURE 32. Styringomyia simplex, male specimen from Brazil (Mato Grosso, Diamantino). Three-dimensional structure of terminalia, dorsolateral view. Abbreviations: aed, aedeagus; aed sh, aedeagal sheath; goncx, gonocoxite; gonst, gonostylus; S9, ninth sternite, vb, ventral branches of aedeagal sheath.

DESCRIPTION. Dimensions. See Table I. Morphology. Ninth sternite (Fig. 32): abruptly narrowed at apex. Tenth tergite (Fig. 22): posterior margin trilobulate; medial

NEOTROPICAL STYRINGOMYIA © 2003 Magnolia Press 21 ZOOTAXA lobe triangular, broader than long. Gonocoxite (Figs. 31-32): distal half conical, abruptly 253 narrowed; longest and more stout hairs along the distal parts of dorsal margin to apex; other hairs slender, variable in length, the longest along the lateral margin. Gonostylus (Figs. 14, 31-32): more or less S-shaped, bearing two strongly sclerotized apical extensions (Fig. 14, III, IV); extension III less developed than extension IV; medial face with a long and stout hair originating near base of extension IV; other hairs of medial face slender and shorter; hairs of lateral face long and slender. Aedeagal sheath (Figs. 31-32): without lateral lobes; dorsal crest four-pointed; space between medial points narrow; apex of each point disposed nearly side by side; ventral branches long, broad at base, abruptly narrowed at tip. Female cercus (Fig. 9): narrowing gradually toward tip; anterior portion of ventral margin interrupted by a constriction.

Styringomyia maya sp. n.

Alexander, 1945b: 241, Figure 8 (male terminalia, identified as S. americana Alexander, 1914).

MATERIAL EXAMINED. Holotype. Male, Ecuador: Azuay, 18 km SE Jesus Maria (2º33S, 79º33W), 1200m, 10 Nov. 1987, J. Rawlins, C. Young, R. Davidson, wet forest habitat (CMNH). Pinned specimen, with dissected terminalia inside microvial with glycerin pinned with the specimen. Paratypes. 2 males and 2 females, same data of holotype (CMNH). Additional material. 1 male, Ecuador, Manabi, Palmar (2º00S, 80º21W),200 m, May 20, 1941, David Laddey (USNM). 1 male, Honduras, Tela, Lancetilla (15º41N, 87º26W), IX-17-1953 (USNM); 1 male, Costa Rica, Rivas, Talamanca (9º25N, 83º37W), January, 1939 (name of collector not legible, USNM); 1 male, British Honduras (Belize), Orange Walk (18º06N, 88º31W), October 18, 1925, (name of collector not legible, USNM); 1 male, Sevilla, Colombia, S. A., (4º16N, 75º58W) V-7-1926, F. W. Walker (USNM); 1 male, Panama, Pina (8º4.2N, 78º11W), January 14, 1954, Field (USNM). ETYMOLOGY. This species is named after the Maya civilization, which in Pre- Colombian times occupied areas that include the most northern parts of the range of its distribution. DESCRIPTION. Dimensions. See Table I. Morphology. Ninth sternite (Fig. 34): abruptly narrowed at apex. Tenth tergite (Fig. 23): posterior margin trilobulate; medial lobe subtriangular, broader than long. Gonocoxite (Figs. 33-34): distal half bifurcated; dorsal branch bearing a tuft of stout hairs, the three longest at its posterior margin; ventral branch with one long and stout hair at apex; other hairs long and slender. Gonostylus (Figs. 15, 33-34): with two strongly sclerotized apical extensions (Fig. 15, III and IV), the apicalmost (extension IV) slightly longer and broader than the other; posterior margin constricted at base, dilated at apex; thin hairs spread along most distal parts of lateral face. Aedeagal sheath (Figs. 33-34): without lateral lobes; dorsal crest two-pointed, with points

22 © 2003 Magnolia Press RIBEIRO disposed side by side; ventral branches long, broad at base, abruptly narrowed at tip. ZOOTAXA Female cercus: narrowing gradually toward tip; ventral margin gently curved, not con- 253 stricted. REMARKS. Charles Paul Alexander insisted in associating specimens of this species with S. americana (e. g. Alexander, 1945b: 241 (Figure 8), 264; Alexander, 1947: 356 (List of Species)), without, however, clarifying the grounds on which these associations were based. The examined specimens of this new species deposited at USNM were all identified by Alexander as belonging to S. americana.

FIGURE 33. Styringomyia maya sp. n., male specimen from Ecuador (Manabi, Palmar). Aspect of part of terminalia. Specimen on slide. Abbreviations: aed, aedeagus, goncx, gonocoxite; gonst, gonostylus.

FIGURE 34. Styringomyia maya sp. n., holotype. Three-dimensional structure of terminalia, dorsolateral view. Abbreviations: aed, aedeagus; goncx, gonocoxite; gonst, gonostylus; S9, ninth sternite, vb, ventral branches of aedeagal sheath.

Styringomyia amazonica sp. n.

MATERIAL EXAMINED. Holotype. Male, Brazil, Amazonas, Tefé, Alvaraes (ca. 3º24S 64º45W) 50m, 24.II.1981, C. Young. (CMNH). Pinned specimen, with dissected terminalia inside microvial with glycerin pinned with the specimen. Paratypes. 2 males plus 10 mating pairs, same data of holotype (CMNH). Additional material. 19 males and 18 females, same data of holotype and paratypes (CMNH); 1 male, Ecuador, Coca (Francisco de Orel- lana, 0º28S, 77º00W), Napo, May 6, 1965, Luiz Peña (USNM); 1 male, Ecuador, Coca, V- 10-1965, Luiz Peña (USNM). ETYMOLOGY. The name of this species refers to its habitat at the Brazilian and Ecuadorian Amazonian Forest. DESCRIPTION. Dimensions. See Table I. Morphology. Ninth sternite (Fig. 36): abruptly narrowed at apex. Tenth tergite (Fig. 24): posterior margin trilobulate, medial lobe triangular, longer than broad. Gonocoxite (Figs. 35-36): distal half abruptly narrowed, rounded at apex; more stout and longest hairs along apical margin; thin and shorter hairs along the most proximal part of medial margin. Gonostylus (Figs. 16, 35-36): a unique

24 © 2003 Magnolia Press RIBEIRO strongly sclerotized extension at apex of posterior margin (Fig. 16, III + IV); anterior mar- ZOOTAXA gin near straight; posterior margin slightly sinuous, rounded at its proximal portion; hairs 253 of lateral face concentrated on a small protuberance near apex. Aedeagal sheath (Fig.36): without lateral lobes; dorsal crest four-pointed, medial points separated by a wide space; ventral branches long, broad at base, narrowing gradually toward tip. Female cercus: narrowing gradually toward tip; ventral margin gently curved, not constricted. REMARKS. The examined specimens of this new species deposited at USNM were identified by Charles Paul Alexander as belonging to S. dorsolineata.

FIGURE 35. Styringomyia amazonica sp. n., male specimen from Ecuador (Francisco de Orel- lana). Aspect of part of terminalia. Specimen on slide. Abbreviations: goncx, gonocoxite; gonst, gonostylus.

FIGURE 36. Styringomyia amazonica sp. n., holotype. Three-dimensional structure of terminalia. Abbreviations: aed, aedeagus; goncx, gonocoxite; gonst, gonostylus; S9, ninth sternite.

Styringomyia manauara sp. n.

MATERIAL EXAMINED. Holotype. Male, Brazil, Amazonas, Manaus (3º06S, 60º00W), 1kmfromTarumafalls,12-II-1981,100m,C.Young(CMNH). Pinned specimen, with dissected terminalia inside microvial with glycerin pinned with the specimen. Paratypes. 1 male and 1 female, same data of holotype (CMNH); 2 males and 2 females, Brazil, Amazo- nas, Manaus (3º06S, 60º00W), 1 km from Taruma falls, 13-II-1981, 100m, C. Young (CMNH ); 1 male, Brazil, Amazonas, Manaus (3º06S, 60º00W), 1 km from Taruma falls, 14-II-1981, 100m, C. Young (CMNH ). Additional material. 1 male and 1 female, Brazil, Amazonas, Paraque, 30 km East. Manaus (3º06S, 60º00W), Amazon R. 18 February 1981, 33.5 m, C. W. Young (CMNH); 1 male, CAMPUS F. U. A. Manaus (3º06S, 60º00W), Am., BRASIL, Arm. ESALQ Blb-Bl, Data: 08-09/XI/82, col. P. F. Buhenheim (INPA). ETYMOLOGY. From the Tupi-Guarani indigenous language, relative to or coming from Manaus.

26 © 2003 Magnolia Press RIBEIRO DESCRIPTION. Dimensions.SeeTableI.Morphology. Ninth sternite (Fig. 37): nar- ZOOTAXA rowing gradually toward tip. Tenth tergite (Fig. 25): posterior margin slightly trilobulate; 253 medial lobe rounded, much broader than long. Gonocoxite (Fig. 37): distal half abruptly narrowed, with a stripe of stout hairs all along its laterodorsal margin to apex; other hairs longer and slender. Gonostylus (Figs. 17, 37): long and narrow, curved, with three strongly sclerotized apical extensions; extensions III and IV closely jointed; extension II more separated. Aedeagal sheath (Fig. 37): without lateral lobes; dorsal crest three-pointed, medial point anterior in relation to the others; ventral branches long, narrowing gradually toward tip. Female cercus: narrowing abruptly toward tip; ventral margin gently curved, not constricted.

FIGURE 37. Styringomyia manauara sp. n., male paratype. Three-dimensional structure of terminalia. Abbreviations: aed, aedeagus; goncx, gonocoxite; gonst, gonostylus; S9, ninth sternite; T10, tenth tergite.

Phylogeny

Twenty characters of the morphology of the male terminalia were scored for the eight rec- ognizable Neotropical species of Styringomyia (no diagnostic nor phylogenetic informative characters could be scored for S. americana Alexander and S. dorsolineata Alexander, which are treated in this review as nomina dubia) plus four out-group species of Styringo- myia from different biogeographic regions. The data matrix is shown in Table III. Informa- tion on the species used as out-groups is furnished in the Appendix. Character delimitation

NEOTROPICAL STYRINGOMYIA © 2003 Magnolia Press 27 ZOOTAXA and coding followed the conventional approach (sensu Hawkins et al., 1997 and Hawkins, 253 2000), in which the underlying concept behind primary homology assessment is that of the transformational homology, i. e., character states are explicitly considered as transforma- tions. Except for character 20, for which an intermediate condition between its three states is observed, all characters are treated as unordered. Character descriptions are summarized in Table II.

Table II. List of 20 characters scored for the eight Neotropical species of Styringomyia included in the phylogenetic analysis. The weights applied by successive weighting are given between square brackets after each character description.

Ninth sternite: 1- Narrowing gradually toward tip (0); narrowing abruptly toward tip (1). [0.40] Outer gonostylus: 2- absent (0); present (1). [1.00] Aedeagal sheath: 3- lateral lobes: absent (0); present (1). [0.37] 4- ventral branches: absent (0); present (1). [1.00] 5- ventral branches: long (0); short (1). [0.33] 6-medialpointofdorsalcrest:absent(0);present(1).[1.00] 7- lateral points of dorsal crest: simple (0); bifid (1). [1.00] 8- lateral points of dorsal crest: widely separated: (0); separated by a narrow space (1). [0.00] Gonocoxite: 9- apical hairs: long (0); short (1). [1.00] 10- distal part: simple (0); bifurcated (1). [0.00] 11- small lobe bearing curved and stout hair near apex of ventral margin: absent (0); present (1). [1.00] Inner gonostylus: 12- much longer than broad (0); almost as long as broad (1). [1.00] 13- spine-like hairs: absent (0); present (1). [0.33] 14- spine like hairs: short (0); long (1). [1.00] 15- extension I: absent (0); present (1). [1.00] 16- extension II: absent (0); present (1). [1.00] 17- extension IV: absent (0); present (1). [1.00] Tenth tergite: 18- posterior margin: rounded (0); trilobulate (1); V-shaped (2). [0.53] 19- incision of posterior margin: tenuous (0); deep (1). [1.00] 20- medial lobe of posterior margin: rounded (0); slightly triangular (1); noticeably triangular (2). [1.00]

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 S. maya sp. n. 1001000101000-001111 S. amazonica sp. n 1001001000000-001112 S. simplex 1001001100000-001112 S. manauara sp. n. 0001010000000-011100 S. mystica 001101000010111110- - S. youngi sp. n. 001101000110111100- - S. atlantica sp. n. 00111???1001101112- - S. paulista 00111???100110111100 S .annulipes 11011?00000010***110 S. crassicosta 1110-100000010***0- - S. sp. 1 1100-100000010***0- - S. sp. 2 01011100010010***0- - S. sp. 3 1100-100010010***0- -

The morphology of the inner gonostylus of the Neotropical species of Styringomyia differs completely from that of the species from other regions, and for a set of characters relating to its morphology (characters 15-17), no comparison could me made with any out- group taxa. For this reason they were all treated as non-comparable and coded as missing data. The analysis of the data matrix with all characters receiving equal weights yielded 4 most parsimonious cladograms with 31 steps, CI = 0.70, and RI = 0.80. The relationships between the in-group taxa is constant in all the most parsimonious trees obtained. A posteriori successive weighting based on the rescaled consistency index yielded a single, fully resolved cladogram with Length =18.48, CI = 0.89, and RI = 0.93, in which the topology within the in-group (Fig. 38) is the same obtained in all the most parsimonious cladograms of the former analysis. Because it is intended to furnish a phylogenetic hypothesis on which the less reliable characters have the minimal possible influence on the pattern given by the relatively more congruent characters, the results of the analysis applying aposteri- ori weighting is the one considered for discussion. The distribution of the characters along the proposed phylogeny is discussed below, following the clades indicated by letters in Fig. 38. Only non-autapomorphic characters providing unambiguous support for clades are plotted in Fig. 38, although reverence to the other characters are also made along the text. Clade A. The monophyly of the clade including all Neotropical species of Styringo- myia is supported in the present study by the loss of the outer gonostylus (character 2, state 0). The presence of the outer gonostylus is probably plesiomorphic for the Tipulomorpha as a hole. In Styringomyia, this loss is known to occur also only in the species S. scalaris

NEOTROPICAL STYRINGOMYIA © 2003 Magnolia Press 29 ZOOTAXA Alexander, 1960, from Papua New Guinea, but this resemblance with the Neotropical spe- 253 cies may be almost certainly homoplastic. In the out-group species possessing ventral branches in the aedeagal sheath (S. annulipes and S. sp 2), this structure is short. The long ventral branches of the aedeagal sheath (character 5, state 0) is also a synapomorphy unit- ing all the Neotropical species. This character has suffered a reversal in S. paulista and S. atlantica sp. n. (see discussion about Clade E).

FIGURE 38. Phylogenetic relationships of Neotropical Styringomyia species obtained from analysis of the data matrix in Table III. The number refers to the characters and their states (in parenthesis). Autapomorphic and ambiguously optimized characters omitted (see text for details).

30 © 2003 Magnolia Press RIBEIRO Clade B. The ninth sternite narrowing abruptly toward tip is probably plesiomorphic ZOOTAXA for Styringomyia. The ninth sternite narrowing gradually toward tip (character 1, state 0) is 253 a unambiguous synapomorphy supporting Clade B, although it can be found homoplasti- cally in non-Neotropical species of the genus (e. g. in the undetermined S. sp2 of data matrix). The deep incisions of the posterior margin of the tenth tergite are also likely to correspond to the plesiomorphic state, at least at the level of the Neotropical clade. These incisions have become much more tenuous at the base of Clade B (character 19, state 0). All the species of this clade have the extension II of gonostylus. However, due the impossibility of comparison with any out-group species, it is not possible to determine unambiguously whether the presence of extension II (character 16, state 1) of gonostylus is synapomorphic or plesiomorphic at this level. Clade C. The monophyly of Clade C is supported by the presence of lateral lobes in the aedeagal sheath (character 3, state 1) and extension I of gonostylus (character 15, state 1). It must be remarked that lateral lobes in the aedeagal sheath also occurs in other non- Neotropical species of Styringomyia,asinS. crassicosta.

FIGURE 39. Map of South America, showing collecting sites for the Neotropical Styringomyia specimens examined in this work. Points may refer to more than one site in close proximity.

NEOTROPICAL STYRINGOMYIA © 2003 Magnolia Press 31 ZOOTAXA Clade D. Unambiguous support for Clade D is given by the loss of the medial point of 253 the dorsal crest of the aedeagal sheath (character 6, state 0), and the slightly triangular form of the medial lobe of the posterior margin of the tenth tergite (character 20, state1). ACCTRAN optimization puts state 1 of character 8 (a narrow space separating the medial points of the dorsal crest of the aedeagal sheath) as synapomorphic at this level, with a reversal in S. amazonica sp. n.. On the other hand, DELTRAN mode favors the independent appearance of this apomorphic state in S. simplex and S. maya sp. n.. None of these two possible solutions are chosen here for the character. The problem is considered an opened question susceptive to be solved when further evidence become available. In all out-group species and in the species included in Clade C, spine-like hairs in the gonostylus are present (character 13, state 1). This state is probably plesiomorphic for Styringomyia. ACCTRAN optimization puts the absence of spine-like hairs as plesiomorphic for the Neotropical clade, with a reversal at the base of Clade C. This hypothesis is considered here as improbable. DELTRAN mode of optimization is chosen here for this character, which points for the independent loss of the spine-like hairs of gonostylus (character 13, state 0) at the base of Clade D and in S. manauara sp. n.. Clade E . The sister group relationships between S. paulista and S. atlantica sp. n. is well supported by short ventral branches of the aedeagal sheath (character 5, state 1), short hairs at apex of gonocoxite (character 9, state 1), and gonostylus almost as long as broad (character 12, state 1). There are other similarities between these species, relating mainly with the morphology of the gonostylus (compare Figs. 12 and 13), but due to the impossi- bilities of comparison with out-groups, it was not possible in this study to determinate what of such similarities also correspond to synapomorphies. In S. atlantica sp. n., the posterior margin of the tenth tergite changed from the plesiomorphic trilobulate condition to V-shaped (character 18, state 2). The states of characters 6, 7 and 8 could not be scored in the available specimens of the species included in Clade E. Clade F. The hypothesis of S. mystica and S. youngi sp.n.assistergroupsisunambig- uously supported by the presence of a small lobe bearing a curved stout hair near the apex of the ventral margin of gonocoxite (character 11, state 1; Figs. 26-28, arrow), long spine- like hairs in the gonostylus (character 14, state 1), and the rounded form of the posterior margin of the tenth tergite (character 18, state 0). As is the case for the pair of species in Clade E, the impossibility of comparison with out-groups makes difficult to determine which of the remarkable similarities in the morphology of the gonostylus in S. mystica and S. youngi sp. n. (Figs. 10 and 11) also correspond to synapomorphies. Clade G. The bifid lateral points of the dorsal crest of the aedeagal sheath (character 7, state 1) and the well defined triangular form of the medial lobe of the tenth tergite (character 20, state 2) support the sister group relationship between S. amazonica sp.n.andS. simplex.

32 © 2003 Magnolia Press RIBEIRO Acknowledgements ZOOTAXA 253 I am very indebted to Dr Dalton de Souza Amorim (Universidade de São Paulo) for the advisement in the preparation of this work. Special thanks to Dr Conrad Labandeira (Smithsonian Institution) for hosting me as a visitor in the National Museum of Natural History in Washington, DC in December of 2001. I would like to thank Dr Wayne Mathis, curator of Diptera of the National Museum of Natural History for allowing the study of the collection under his care. I am indebted to Holly Williams (Smithsonian Institution), for the assistance in accessing the specimens in the Alexander Collection of Crane flies and to Finnegan Marsh (Smithsonian Institution) for assistance in taking the photographs and other facilities. For the loan of the holotype of Styringomyia paulista IamindebtedtoDr Eliana Marques Cancello (Museu de Zoologia da Universidade de São Paulo, Brazil). I thank Dr Chen Young (Carnegie Museum of Natural History, USA) and Dr John Gelhaus (The Academy of Natural Sciences, Philadelphia, USA) for the prompt support in sending on loan the Styringomyia specimens from the collections under their care. I am grateful to Dr Sigitas Podenas (Vilnius University, Lithuania), Dr. Vladimir Blagoderov (American Museum of Natural History, New York, USA) and two anonymous reviewers for the cor- rections, critics and suggestions that greatly improved the quality of the final version of this paper. Most of this research was financially supported by a Master Fellowship by FAPESP (Research Foundation of the São Paulo State, Brazil; process 2001/02703-1). During the final stage of preparation of this paper I was financially supported by a Ph.D. fellowship by CNPq (Brazilian National Research Council).

References

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34 © 2003 Magnolia Press RIBEIRO APPENDIX. Information on the species used as out-groups in the phylogenetic analysis. ZOOTAXA 253 DETERMINED MATERIAL Styringomyia annulipes Enderlein, 1912. CAMEROON: Southwest Prov. Bakingili 25 km W. of Limbe 10m 1-10 June 1984 J. Rawlins, R. Davidson, J. DiGuilio. det.: G. C. Ribeiro (CMNH). Styringomyia crassicosta Speiser, 1908. CAMEROON: Southwest Prov. Bakingili 25 km W. of Limbe 10m 1-10 June 1984 J Rawlins, R. Davidson, J. DiGuilio. det.: G. C. Ribeiro (CMNH).

UNDETERMINED MATERIAL Styringomyia sp 1. TAIWAN: Taitung. Chihpen. 70m 16 March 1988 C. Young, J. Rawlins. (CMNH). Styringomyia sp. 2. INDONESIA: Sulawesi Utara Dumoga-Bone N. P. 10 Aug. 1985 Coll. Chen Young. (CMNH). Styringomyia sp. 3. MALAWI: Chitipa District Mughese Forest reserve 09-39S 33-32E 1890m 28 January 1989 J. Rawlins, S. Thompson. (CMNH).

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