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Studies on Neotropical Fauna and Environment 2004, Vol. 39, No. 1, pp. 15–35

New species of Fittkau, a Member of the -Group (Diptera: : ), with a Review of the

Humberto F. Mendes1, Trond Andersen2 and Ole A. Sæther2

1Department of Biology-FFCLRP, University of São Paulo, Ribeirão Preto, Brazil 2Museum of Zoology, University of Bergen, Bergen, Norway

Abstract Emended diagnoses of all stages of the genus Ichthyocladius Fittkau (1974) mentioned several larvae of the genus Fittkau are given. The genus appears to be a basal member from Trinidad, Venezuela, Guyana, Bolivia and Brazil not of the Corynoneura group showing affinities also to the Euki- identified to species. In the present paper all stages of two efferiella group. A number of features apparently are the new Brazilian species and the pupal exuviae of two addi- result of the phoretic life on catfishes. Ichthyocladius kro- tional species from Brazil and Argentina are described and nichticola sp. n. and I. lilianae sp. n. are described and figured. figured in all stages. Additional pupae from the Amazonas in Brazil and from the Parque Nacional Iguazú in Argentina are also described and figured. Keys to pupae and larvae of the Material and methods genus are given. The general terminology follows Sæther (1980). The male Keywords: Chironomidae, Orthocladiinae, Corynoneura – hypopygium is figured with tergite IX removed, dorsal group, Ichthyocladius, new species, Brazil, Argentina. aspect to the left, and ventral aspect to the right. Dorsal view of tergite IX, gonocoxite and gonostylus are figured separately. Introduction All material is mounted on microscope slides and prepared in Canada Balsam or Euparal. The holotypes Fittkau (1974) erected the genus Ichthyocladius for two and paratypes of the new species are deposited at the species from Peru and Ecuador. The larvae of the genus live Museu de Zoologia da Universidade de São Paulo, Brazil phoretically on catfishes of the families Astroblepidae and (MZUSP). Paratypes are also deposited at the Zoologische , attaching themselves to the skin and scales of Staatssammlung, Munich, Germany (ZSM), and at the the fish. At the end of the fourth larval stage the larvae spin Museum of Zoology, University of Bergen, Norway a holdfast that is attached to spines or fin rays and finally (ZMBN). formed into a pupal case. The only named species, I. neotrop- icus Fittkau, was described based on pharate pupae and asso- ciated larvae. Several details, especially concerning the Ichthyocladius Fittkau wings and the apodemes of the male hypopygium, thus have not been described previously. Ichthyocladius Fittkau 1974: 91.

Received: 19 January 2003 Accepted: 21 November 2003 Correspondence: T. Andersen, Museum of Zoology, University of Bergen, Muséplass 3, N-5007 Bergen, Norway. Fax: +47 55589677; E-mail: [email protected]

DOI: 10.1080/01650520412331270936 © 2004 Taylor & Francis Ltd. NNFE391015 8/17/04 2:13 PM Page 16

16 H.F. Mendes et al.

Type species and some apical ventrolateral setae; laterosternite IX bare. Sternapodeme strong, triangular. Phallapodeme and aedeagal Ichthyocladius neotropicus Fittkau, 1974, by original desig- lobe normally developed. Virga absent. Gonocoxite well nation and monotypy. developed, superior volsella apparently absent, inferior volsella low to moderately well developed. Gonostylus Other included species broadest near apex, without crista dorsalis and megaseta. Female genitalia with well developed gonocoxite with Ichthyocladius kronichticola sp. n.; Ichthyocladius lilianae several setae. Tergite IX very large, covering most of geni- sp. n.; Ichthyocladius sp. Ecuador; Ichthyocladius sp. Rio talia, medially divided, with several apical setae. Segment X Marauiá, Brazil; Ichthyocladius sp. Argentina. apparently normal. Postgenital plate weak or absent. Cercus triangular, small. Gonapophysis VIII undivided. Shape of apodeme lobe and coxosternapodeme unknown. Seminal Diagnosis capsules much larger than cerci, ovoid; with orally placed Imago. Small species, wing length about 1.3mm. neck. Spermathecal ducts nearly straight, apparently with Eye bare, rounded, no dorsomedian elongation. Antenna common opening. Labia not known. with 12 (or perhaps sometimes fewer, see below) flagellom- eres in male, six in female; male antenna sparsely plumed; Pupa. Small pupae, about 2–3mm long. Exuviae nearly groove beginning on flagellomere 3; sensilla chaetica minute, transparent to strongly marked with brown. present on flagellomeres 2, 3 and 12; apex without straight Cephalic tubercles and frontal warts absent. A low protu- apical seta; AR lower than 0.4. Only one palpomere, no sen- berance densely covered with tubercles present basally on silla clavata. Temporals consisting of a few minute inner and pedicel. Frontal setae absent, frontal apotome wrinkled. Tho- outer verticals (or sometimes absent?). Tentorium with broad racic horn absent. Wing sheath smooth, without pearls or but short base, stipes apparently normal. Cibarial pump nose, but sometimes with notch. Thorax with median tuber- broad, but short, with concave anterior margin and well cle. Three precorneals, four antepronotals, at least one pos- developed, triangular cornua. Clypeus narrow with short torbital, and four dorsocentrals present, all minute. lateral setae. Segment I and sternite IX without shagreen; tergites and Antepronotal lobes broad, collar-like, not medially nar- sternites II–VIII with shagreen, on tergites II–VII median rowed, without setae. Scutal tubercle present and prominent. shagreen spinules stronger and sometimes placed on protu- Acrostichals absent, dorsocentrals uni-triserial, several pre- berance. Conjunctives bare. Tergites II–V and sternites alars, supraalar present. Scutellum with setae transversely IV–VII each with 34–50 caudal hooklets; a few lateral or uniserial. median hooklets sometimes present also on tergite I and/or Wing broad, with well developed, but not projecting anal sternite III. Pedes spurii A and B absent. lobe; membrane with distinct punctation; costa slightly Abdominal setae minute; segments I–V each with 1L extended; veins R and R short, thick and fused with costa 1 2+3 seta, VI–VII with 1–2L setae, VIII with four L setae; tergites in thick clavus, ending slightly beyond midpoint of wing; I–VI with four, VII with three and VIII with one pair of D R weak; VR moderately high; Cu slightly sinuous; postcu- 4+5 1 setae; sternite I with three, II with four, III–VII with five, and bitus ending distal to cubital fork, anal vein ending below or VIII with two pairs of V setae. Segment IX with three short, slightly distal to cubital fork. Brachiolum with one seta, other scimitar-like or hair-like curved macrosetae situated ven- veins bare. Squama bare. Sensilla campaniformia about eight trally or at apex. O setae not observed. Genital sac of male basally and eight apically on brachiolum, apparently none slightly to much longer than anal segment, broadly rounded, below setae on brachiolum; one present basally on subcosta, without shagreen or apically wrinkled and with a few minute and one on RM. apical spinules. Front leg ratio high, about 0.8. Front tibia with straight, weak spine-like tibial spur; spurs of mid tibia either with thorn-like lateral denticles or entirely consisting of thorn-like . Small larvae, up to 7mm long. spines; hind tibia with well developed spurs covered with Antenna with four segments, basal segment about as long thorn-like lateral denticles; hind tibial comb absent or as or slightly longer than flagellum, second antennal segment reduced to a few setae. Short tarsal pseudospurs present on slightly longer than or subequal to third. Basal antennal

metatarsi of mid and hind legs and ta2 of mid leg. Pulvilli segment 1.5–2 times as long as basally wide, with ring organ absent. No sensilla chaetica observed. situated in apical third. Lauterborn organs not observed, style Tergites II–VII in male, II–VIII in female each with group well developed. Blade slightly shorter than flagellum, acces- of median setae sometimes on a low protuberance, tergites sory blade about half as long. except for male tergite IX otherwise bare; sternites except for S I pectinate to plumose. Other S setae simple. Labral female sternite VIII bare. lamella large, bluntly triangular with about 50 teeth. Chaetae Tergite IX of male large, covering much of gonocoxites, reduced to two pegs. Pecten epipharyngis consisting of three with notched or bilobed posterior margin, with several apical pointed teeth. About six chaetulae laterales with the median NNFE391015 8/17/04 2:13 PM Page 17

New Species of Ichthyocladius 17

apparently serrated, chaetulae basales weak. Premandible Characters in common for all the genera of the two groups with three distinct dark inner teeth and 3–5 broadly lamel- include absence of pedes spurii A and B of the pupa; sex late, pale and fused outer teeth (not visible in all views), dimorphism of sternite VIII of the pupa; gonapophysis VIII without or with vestigial brush. of the female genitalia undivided; and ventromental plates of Mandible with long, bent apical tooth longer than com- the larvae narrow, inconspicuous, widest posteriorly and bined width of 3–4 inner teeth. Seta subdentalis well devel- without setae underneath. oped. Seta interna absent. Ichthyocladius, in addition to the possession of a clavus, Mentum with 16–20 equal-sized small median teeth shares the reduced temporals, the large tergite IX covering nearly in a straight line and five pairs of slightly larger lateral most of the male hypopygium, absence of anal point and teeth. Ventromental plates long and narrow. Setae submenti virga, seminal capsule with orally placed neck, and situated above base of outer lateral tooth of mentum. nearly straight spermathecal duct with Corynoneura Maxilla relatively large, pecten galearis apparently absent, and Kieffer. An inverted V-shaped ster- maxillary palp well developed. Anterior lacinial chaeta napodeme without oral projection is present also in apparently with apical teeth. Corynoneura. However, Ichthyocladius also has several fea- Anterior and posterior parapods well developed; claws of tures not found in Corynoneura and Thienemanniella. Some anterior parapods pale to brown, longer claws smooth or with of these, such as the reduced larval and pupal features and a few very small inner teeth; posterior parapods with 12–15 the absence of a megaseta on the male gonostylus, likely are black claws without teeth. Procercus absent or very low, 3–4 a result of the phoretic life style, while others appear to be short anal setae. Supraanal seta minute. Anal tubules very plesiomorphies. In the last group are non-cordiform tarsi, short. Body setae very short. presence of tarsal pseudospurs, and the well developed anal lobe and distinct punctations of the wings. That the clavus ends in the distal half of the wing instead of in the basal half Diagnostic characters also appears to be a plesiomorphy as compared to other The male imagines are separable from all other chironomids genera of the Corynoneura group (Sæther & Kristoffersen, by the presence of a clavus combined with a bare scutal 1996). tubercle and the lack of a gonostylar megaseta. The pupae Other characteristic features include the scutal tubercle differ from all chironomids by the possession of hooklets on without any microtrichial tuft or acrostichals which other- tergites I–IV and sternites IV–VII combined with lack of a wise amongst the Orthocladiinae occurs only in thoracic horn and curved (associated pupae) or hair-like Brundin. The one-segmented palp otherwise is found only in (un-associated pupae) macrosetae mostly on the ventral side some species of Symbiocladius Kieffer (Gonser & Spies, of the anal lobe. The larva differs from other chironomids 1997). At first the pupa was thought to lack setae, spines or by having about 16–20 dark, equal-sized median and five thorns from the pupal anal segment as otherwise found only pairs of slightly larger lateral teeth in the mentum and no in very few genera or species including Rheosmittia and mandibular seta interna. ancyla Svensson, a commensal in Ancylus flu- viatilis Müller (Svensson, 1986). However, there are three macrosetae mostly placed ventrally on the anal lobe between Systematics the lobe and the genital sac and a minute additional apico- The presence of a clavus places the genus near Corynoneura lateral seta. These setae may easily have been overlooked also Winnertz and related genera, while the caudal hooklets of the in other genera. Fittkau (1974: fig. 11) is incorrect in describ- pupa combined with the absence of a fringe on the anal lobe ing the S setae, S I really is S II and S II is S III, while S I indicate relationship with the Eukiefferiella () is figured as part of the pecten epipharyngis. Thus S I is in group. However, the two groups may be sister groups. fact pectinate or plumose, a feature found in Kieffer The presence of large caudal hooklets on the tergal con- of the Eukiefferiella group, but not in genera near junctives or posterior on the tergites is conspicuous both in Corynoneura. The labral lamella is large, bluntly triangular Ichthyocladius, Tempisquitoneura Epler and a new genus and finely pectinate or denticulate, but often is perpendicu- near Thienemanniella Kieffer and every bit as conspicuous lar to the labral surface and thus obscured. A large labral as those in the Eukiefferiella group. Sæther (1977) placed the lamella is not found in any genus near Corynoneura or Euki- Corynoneura group as sister group to a reduced Orthocladi- efferiella. S I appear to be attached underneath the labral ini sensu Brundin and a Metriocnemini sensu Brundin com- lamella, i.e., about as in v. d. Wulp (Cranston bined, i.e., with some plesiomorphic groups removed, with et al., 1983: fig. 9.43). Spärck as sister group to all three groups The larval mentum is of the same type as the mentum of combined and the Eukiefferiella group as sister to all the some other phoretic larvae such as Baeoctenus Sæther, Drat- above. That is, the Corynoneura group was just two nodes nalia Sæther et Halvorsen and Zavrˇel. Sæther removed from the Eukiefferiella group. The new genus near (1986) discuss the menta of these genera as a prime example Thienemanniella, Tempisquitoneura and Ichthyocladius, of parallel selection and suggests that the menta act as combs however, strengthen the indications of relationship. scraping off particles from hairs and appendages of their NNFE391015 8/17/04 2:13 PM Page 18

18 H.F. Mendes et al.

hosts. Of the above genera, Dratnalia belongs in the Eukief- pharate male appears to be lost. Among other things it thus feriella group. Amongst genera related to Corynoneura, Tem- is not clear how many flagellomeres are present. Fittkau pisquitoneura is known to be symphoretic on Corydalus (1974) mentions that the male antenna is as in the female, (Megaloptera: Corydalidae) (Epler & De la Rosa, 1995). which could mean that there are six flagellomeres only. Tem- However, in that genus the immatures are normal. porals also are mentioned to be absent. However, the tempo- ral setae present in the new species described here are tiny and could easily be overlooked. Key to known pupae of Ichthyocladius Fittkau Fittkau (1974) also mentions that the claws of the anterior 1. Tergites II–VII with strong median shagreen spinules parapods are pale, thus contrasting with the black claws of placed on protuberance. Brazil (Figs. 11–15) ...... the posterior parapods. In the examined larvae, however, the ...... I. kronichticola sp. n. claws of the anterior parapods are brown although not as dark – Tergites II–VII without protuberance or protuberance at as in the new species described below. most indicated on tergite VI and VII ...... 2 Fittkau overlooked the anal macrosetae of the pupae. In 2. Tergite I without posterior spines or hooklets, hooklets addition to three ventromedian, strongly curved, slender on tergite II distinctly weaker than those on tergites macrosetae, slightly widened in apical third, there is one III–IV, exuviae pale. Peru (Fittkau, 1974: figs. 4 & minute apical seta as also observed in most other Ichtyocla- 5) ...... I. neotropicus Fittkau dius exuviae. – Tergite I with weak posterior hooklets, hooklets on tergite II as strong as those on tergites II–IV ...... 3 3. Exuviae pale; anal macrosetae strongly curved, scimitar- Ichthyocladius kronichticola sp. n. (Figs. 1–20) like, broad. Brazil (Figs. 23–25) ...... I. lilianae sp. n. Type material – Exuviae amber with dark brown margins; anal macrosetae hair-like ...... 4 Holotype male with larval and pupal exuviae: Brazil, São 4. Abdominal apex rounded, gradually narrowed between Paulo State, Parque Estadual Intevales, Iporanga, Rio da segment VIII and IX, female genital sac ending far short of Mortes, 17.VIII.2000, H.F. Mendes (MZUSP). Paratypes: apex of anal segment. Brazil (Figs. 32, 33) ...... one male, Brazil, São Paulo State, Iporanga, Petar, ...... I. sp. Rio Marauiá 24°33¢52≤S 48°40¢35≤W, 3.IV.1997, S. Buck (ZSM); five – Abdomen abruptly narrowed between segment VIII and larvae, São Paulo State, Parque Estadual Intervales, Ipo- IX, male genital sac much longer than anal segment and con- ranga, Poços Altos, 1.II.2000, P.C. Bispo, A.S. Melo & V.R. spicuously bicolored, female genital sac reaching apex of de Ribeiro (MZUSP, ZMBN, ZSM). anal segment. Argentina (Figs. 34, 35) . . . . I. sp. Argentina

Etymology Key to known larvae of Ichthyocladius Fittkau From Kronichthys, the genus of Loricariid fishes on which 1. Mandible with three inner teeth. Ecuador (Fittkau, 1974: the larvae where situated, and – cola, Latin suffix meaning fig. 16) ...... I. sp. Ecuador inhabiting. Regarded as a noun in apposition. – Mandible with four inner teeth ...... 2 2. Procercus absent, three anal setae. (Fittkau, 1974: fig. 13) ...... I. neotropicus Fittkau Description – Procercus present, four anal setae ...... 3 Male imago (n = 2, except when otherwise stated). 3. Mentum with 16–18 median teeth and five pairs of lateral Total length 2.02–2.30mm. Wing length 1.32–1.34mm long. teeth (Figs. 16–20) ...... I. kronichticola sp. n. Total length/wing length 1.51–1.80. Wing length/length of – Mentum with 20 median teeth and five pairs of lateral teeth profemur 3.29–3.52. Thorax and head dark brown; abdomen (Figs. 26–31) ...... I. lilianae sp. n. dark brown with posterior 1/4 of segment VI and posterior half of VII pale; coxa, trochanter, femora and tibia dark brown with setae on fore tibia in pale spots, tarsi pale; wings Ichthyocladius neotropicus Fittkau dark. Ichthyocladius neotropicus Fittkau, 1974: 101. Head (Fig. 1). Antenna (Fig. 3) with 12 flagellomeres, AR 0.21–0.28. Ultimate flagellomere 100–136mm long. Tempo- Material examined ral setae consisting of three minute outer verticals and about Paratypes: Peru, Rio Tambopata, tributary of upper Rio five minute inner verticals. Clypeus with 9–13 setae. Tento- Madeira, one male pupa, one pharate female pupa, one larva, rium (Fig. 2) 135mm long, 38 wide. Palpomere 43–45mm 5–6.XI.1951, H. Heller (ZSM). long. The species is described in detail by Fittkau (1974). Thorax (Fig. 4). Dorsocentrals 14–19, in 1–3 rows; pre- However, his adult specimens were pharate and the illustrated alars 4–7. Scutellum with nine setae. NNFE391015 8/17/04 2:14 PM Page 19

New Species of Ichthyocladius 19

Figs. 1–10. Ichthyocladius kronichticola sp. n., male imago: (1) Head. (2) Tentorium and cibarial pump. (3) Antennal apex. (4) Thorax. (5) Apex of front tibia. (6) Apex of mid tibia. (7) Apex of hind tibia. (8) Wing. (9) Tergite IX and dorsal aspect of gonocoxite and gonostylus. (10) Hypopygium with tergite IX and gonostylus removed: left dorsal aspect, right ventral aspect.

Wing (Fig. 8). VR 1.18–1.20. Clavus length 735–776mm Hypopygium (Figs. 9, 10). Tergite IX with 14–17 long. Clavus length/wing length 0.55–0.56. Cu/Wing length apical dorsal setae and 12–14 apicoventral setae. Phal- 0.61 (1). lapodeme 60mm long. Gonocoxite 184mm long, inferior Legs (Table 1). Spur of front tibia (Fig. 5) 21mm long; volsella moderately developed; gonostylus 82mm long, spurs of mid tibia (Fig. 6) 21–23 and 37–41mm long; spurs with a few weak setae at apex, longest 5–7mm long. HR 2.25; of hind tibia (Fig. 7) 18 and 21(1)mm long. No comb setae. HV 2.46–2.80. Pseudospurs each about 10mm long. NNFE391015 8/17/04 2:14 PM Page 20

20 H.F. Mendes et al.

Figs. 11–20. Ichthyocladius kronichticola sp. n., pupa (Figs. 11–15) and larva (Figs. 16–20). (11) Tergites. (12) Sternites. (13) Frontal apotome and antennal sheath. (14) Thorax and wing sheath. (15) Macrosetae. (16) Labrum. (17) Mentum. (18) Mandible. (19) Antenna. (20) Apex of abdomen.

Table 1. Lengths (in mm, n = 1–2) and proportions of the legs of Ichthyocladius kronichticola sp. n.

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR

P1 380–400 360–380 300–312 120 64–68 32 32–40 0.79–0.86 4.14–4.25 2.44–2.53 2.5–2.7

P2 360 440–460 260–280 128 72 32 40 0.56–0.64 3.97 2.86–3.16 1.9

P3 528 508 328 144 80 32 28 0.65 4.80 3.16 2.4 NNFE391015 8/17/04 2:14 PM Page 21

New Species of Ichthyocladius 21

Pupa (n = 1). Total length 2.79mm. Exuviae dark amber with Diagnostic characters rows of very dark hooklets, genital sacs darker, abdominal The male imago of I. kronichticola sp. n. differs from that of segments darker in median and lateral parts. I. neotropicus by having a larger inferior volsella, presence Cephalothorax. Frontal apotome and antennal sheath as in of outer verticals and 12 flagellomeres. The pupa differs from Fig. 13. Thoracic setae (Fig. 14) less than 50 m long, m other species by having strong median spinules placed on a Dc –Dc 34 m, Dc –Dc 60 m, Dc –Dc 69 m. 1 2 m 2 3 m 3 4 m protuberance on tergites II–VII. The larva has a relatively Abdomen (Figs. 11, 12). Tergite (T) II–VII with strong larger seta subdentalis, and a small, but distinct procercus median shagreen spinules placed on protuberance, T VIII and with four anal setae. IX with strong median shagreen, genital sac apically wrin- kled and with a few minute spinules. Sternites (S) I and II with weak shagreen, S III–VIII with strong shagreen. Sha- Biology green on sternites weaker than on tergites. T I with 33 strongly reduced caudal hooklets or spines of which two are The larvae were taken on Kronichthys sp. Most of the host distinct, dark hooklets; number of subequal, stronger hook- fishes were collected under decayed leaves along the banks lets on T II–V as 41, 46, 37, 33; on S IV–VII as 42, 39, 38, of a fourth order stream, where the water flowed very slowly 32. Macrosetae (Fig. 15) scimitar-like, slender in basal 2/3 (Melo & Froehlich, 2001, stream number 9). The fishes were wide from apical third, tapering to sharp point, about 45mm sampled at 9 to 10 a.m., when they were inactive. A total of long. Additional hair-like seta placed slightly ventral of apex 59 specimens were collected, but only five were carrying about 19mm long. Ichthyocladius larvae. Some other Loricariidae specimens were also collected, three specimens of Neoplecostomus sp. and 12 specimens of Harttia spp., but all without Ichthy- Fourth instar larva (n = 3–4, except when otherwise stated). ocladius larvae. Total length 3.0–3.8mm. Head capsule 0.32–0.34mm long. The fishes were brought to the laboratory individually in Postmentum 126–135mm long. Head light brown, body plastic bags. In order to rear Ichthyocladius adults associated greyish black. with the immatures, each fish carrying only one larva was Head. Antenna as in Fig. 19, antennal segments (in mm): placed in a 15-litre aquarium. The only successful rearing 26–30, 9–11, 8–9, 4–8. AR 1.08–1.33. Basal segment 15–17, was obtained two days after the fishes were collected. 16mm wide; ring organ 21mm (1) from base; blade 24mm (1) Another adult was reared by the same method, but the long; accessory blade not measurable. Labrum as in Fig. 16. exuviae was unfortunately not found. Premandible 50–52mm (2) long. Mandible (Fig. 18) 79– 90mm long. Mentum (Fig. 17) 69–83, 76mm wide; with 16–18 median and five pairs of lateral teeth; distance between Ichthyocladius lilianae sp. n. (Figs. 21–31) setae submenti 36–45, 39 m; ventromental plate 9–15, m Type material 12mm wide. Body. Some claws of anterior parapod with a few Holotype: pharate male pupa with associated larva: weak teeth. All setae on thorax and abdominal segments Brazil, Minas Gerais, Rio Sao Francisco, 20°30¢0≤S I–VIII less than 15mm long. Procercus (Fig. 20) 4–8mm 46°50¢0≤W, 31.XII.1995, L. Casatti (MZUSP). Paratypes: 13 (2) high; 8–9mm wide; with 4 anal setae 49mm (2) long. larvae, same data as holotype except 1993–2000 (MZUSP, Anal tubules 34mm (1) long, 23mm wide at base. Posterior ZMBN). parapods with 13–15 black claws, including 1–3 smaller claws. Etymology

Third instar larva (n = 1–2). Total length 2.22mm. Head Named after Lilian Casatti, who collected the material. capsule 0.23–0.24mm long. Postmentum 90–105mm long. Head light brown, body greyish black. Description Head. Antennal segments (in mm): 15–17, 8, 8–9, 4. AR 0.73–0.82. Basal segment 11–12mm wide, ring organ 11mm Male imago (n = 1). from base, blade 19–21mm long, accessory blade 9mm long. Head. Antenna with 12 flagellomeres, AR 0.28. Ultimate Premandible 33mm (2) long. Mandible 60–64mm long. flagellomere 116mm long. Temporal setae consisting Mentum 26–31mm wide, with 18 median and five pairs of of at least one minute outer vertical and one minute lateral teeth, distance between setae submenti 26–31mm, inner vertical. Clypeus with 13 setae. Palpomere ventromental plate 6–8mm wide. 43–45mm long. Body. All setae on thorax and abdominal segments I–VIII Legs (Table 2). Spur of front tibia 19mm long; spurs of less than 15mm long. Procercus 6mm high, 8mm wide, with hind tibia 19 and 34mm long. No comb setae. four anal setae 45mm long. Posterior parapod 94mm long. Hypopygium (Figs. 21, 22). Tergite IX with altogether 25 Anal tubules 23mm long, 11mm wide at base. setae. Phallapodeme 105mm long, apodeme lobe prominent. NNFE391015 8/17/04 2:14 PM Page 22

22 H.F. Mendes et al.

Figs. 21, 22. Ichthyocladius lilianae sp. n., male imago. (21) Tergite IX and dorsal aspect of gonostylus. (22) Hypopygium with tergite IX and gonostylus removed: left dorsal aspect, right ventral aspect.

Table 2. Lengths (in mm, n = 1) and proportions (approximate) of the legs of Ichthyocladius lilianae sp. n.

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR

P1 431 420 311 109 75 41 43 0.74 4.34 2.94 –

P2 371 334 278 105 64 36 41 0.83 4.00 2.54 –

P3 – – – 120 79 30 43 – – – –

Gonocoxite 176mm long, inferior volsella moderately devel- lateral parts of gula, mentum, mandible and premandible oped; gonostylus 98mm long. HR 1.81. dark brown. Other details not measurable. Head. Antenna as in Fig. 30. Length of antennal segments (in mm): 19–24, 21; 6–9, 7; 5–7, 6; 2–5, 4. AR 1.13–1.43, Pupa (n = 1). Total length 2.79mm. Exuviae pale with 1.27. Basal segment 9–13, 11mm wide; ring organ 11–18, 14 margins of wing sheaths yellowish brown and hooklets mm from base; basal setal mark 5mm from base, distal mark yellowish. 17–19, 18mm (6) from base; blade 12–15, 13mm long; acces- Abdomen (Figs. 23, 24). Tergites (T) II and III with weak sory blade 6–7, 6mm (5) long. Labrum as in Fig. 26. and sparse shagreen, T IV with slightly stronger and more Premandible (Fig. 27) 50–59, 59mm long; with three dark extensive shagreen, T V–IX with strong and extensive sha- inner teeth followed by three broadly lamellate fused teeth, green, genital sac mostly smooth. Sternites (S) I with weak apparently with vestigial brush. Mandible (Fig. 29) 76–104, and sparse shagreen, S II–VIII with stronger and more exten- 84mm long. Mentum (Fig. 28) 73–85, 78mm wide; with 20 sive shagreen. T I with 33 strongly reduced caudal hooklets median and five pairs of lateral teeth; distance between widely interrupted medially; number of subequal, stronger setae submenti 33–45, 41mm; ventromental plate 5–8, 7mm hooklets on T II–V as 36, 37, 34, 36; on S IV–VII as 29, 33, wide. 29, 26. Macrosetae (Fig. 25) about 34mm long, scimitar-like, Body. Some claws of anterior parapod with a few weak broad near base to slightly past middle, then tapering to sharp teeth. All setae on thorax and abdominal segments I–VIII less point; positioned at apex. Additional minute apical seta than 15mm long. Procercus (Fig. 31) 5–7, 6mm (2) high; apparently present lateral of macrosetae. 9–13, 11mm wide; with four anal setae 33–59, 41mm long. Anal tubules 52–71mm (3) long, 19–24mm (3) wide at base. Fourth instar larva (n = 8–11, except when otherwise Posterior parapods with 12 black claws. stated). Total length 1.7–3.0, 2.4mm. Head capsule 0.32–0.36, 0.34mm long. Postmentum 131–165, 145mm Third instar larva (n = 4–5, except when otherwise stated). long. Head light brown with postoccipital margin, postero- Total length 1.53–1.88, 1.69mm. Head capsule 0.23–0.25, NNFE391015 8/17/04 2:14 PM Page 23

New Species of Ichthyocladius 23

Figs. 23–31. Ichthyocladius lilianae sp. n., pupa (Figs. 23–25) and larva (Figs. 26–31). (23) Tergites. (24) Sternites. (25) Macrosetae. (26) Labrum. (27) Premandible. (28) Mentum. (29) Mandible. (30) Antenna. (31) Apex of abdomen.

0.24mm long. Postmentum 90–113, 103mm long. Head light 119–152, 139mm long. Anal tubules 21–28, 24mm long; brown, body greyish black. 11–14, 12mm wide at base. Head. Antennal segments (in mm): 13–17, 15; 6–7, 6; 2–6, 5; 2–4, 4. AR 0.86–1.17, 0.99. Basal segment 7–8, 8mm wide; Diagnostic characters ring organ 7–12, 11mm from base; blade 9–12, 11mm long; accessory blade 6mm (2) long. Mandible 59–73, 65mm long. The male imago of I. lilianae sp. n. differs from that of I. Mentum 40–52, 49mm wide; distance between setae submenti kronichticola sp. n. by having a smaller inferior volsella, 24–36, 29mm; ventromental plate 4–5, 5mm wide. and from I. neotropicus by having more numerous flagel- Body. Procercus 6–7, 7mm high; 7–9, 8mm wide; with lomeres and only two temporals. The males of I. lilianae four anal setae 38–50, 44mm long. Posterior parapod and I. neotropicus, however, are known only from mature NNFE391015 8/17/04 2:14 PM Page 24

24 H.F. Mendes et al.

pupae. The pupa of I. lilianae differs from that of I. Abdomen (Fig. 30). Abdominal apex rounded, gradually kronichticola by lacking strong median shagreen spinules narrowed between segment VIII and IX. Tergite (T) II–VII placed on protuberances on tergites II–VII, from I. neotrop- with median shagreen spinules not on protuberance. T I with icus by having the caudal hooklets of tergite II as strong as 16 weak and small medial hooklets or spines, none dark; those on more posterior tergites, and from the unnamed number of caudal, subequal, stronger hooklets on T II–V as species by having pale exuviae and strongly curved, scimi- 31, 40, 38, 34; on S IV–VII as 30, 23, 32, 26. Macrosetae as tar-like anal macrosetae. The larva has a relatively larger seta in Fig. 33. subdentalis, and a small, but distinct procercus with four anal setae. Ichthyocladius sp. Argentina (Figs. 34, 35) Biology Material examined The larvae were collected on Hypostomus cf. garmani Argentina, Parque Nacional Iguazú, river before the falls, (Regan, 1904) and Harttia sp. three male and one female pupal exuvia, 4.XII.1996, F. Reiss (ZSM). Ichthyocladius sp. Rio Marauiá (Figs. 32, 33) Material examined Description Brazil, Amazonas State, Rio Marauiá, upper half from Pupa (n = 4). Total length 2.65–3.12, 2.79mm. Exuviae dark Cachoeira S. Antônio, surface drift (Abschaum), one female amber with very dark rows of hooklets, genital sacs darker pupal exuviae, 22.I.1963, E.J. Fittkau (ZSM). laterally. Cephalothorax. All thoracic setae less than 50mm long, Distance Dc –Dc 41–94, 77mm; Dc –Dc 8–60, 33mm; Description 1 2 2 3 Dc3–Dc4 124–150, 137mm. Pupa (n = 1). Total length 2.93mm. Exuviae dark amber with Abdomen (Figs. 32). Abdomen abruptly narrowed very dark rows of hooklets, abdominal segments VII and VIII between segment VIII and IX. Tergite (T) II–VII with median darker laterally. shagreen spinules not on protuberance. T I with 27–45, 37 Cephalothorax. All thoracic setae less than 50mm long, weak and small medial hooklets; number of caudal, subequal,

Distance Dc1–Dc2 19mm, Dc2–Dc3 71mm, Dc3–Dc4 81mm. stronger hooklets on T II–V as 34–45, 39; 30–39, 34; 27–43,

Figs. 32–35. Ichthyocladius sp. Rio Marauiá (Figs. 32, 33) and Ichthyocladius sp. Argentina (Figs. 34, 35). (32 & 34) Tergites. (33 & 35) Macrosetae. NNFE391015 8/17/04 2:14 PM Page 25

New Species of Ichthyocladius 25

Table 3. Characters and states used in the phylogenetic analyses.

Imagines 1. Antennal ratio: (0) higher than 0.9; (1) 0.46–0.9; (2) lower than 0.46. 2. Number of flagellomeres: (0) 13; (1) fewer. 3. Male antennal apex: (0) without subapical seta; (1) with. 4. Female antennal apex: (0) without subapical seta; (1) with. 5. Eyes: (0) bare, at most partly pubescent; (1) hairy or pubescent. 6. Palpomeres: (0) five palpomeres of normal length; (1) four or less palpomeres or palpomeres strongly reduced in length. 7. Sensilla clavata of palpomeres: (0) palpomere 3 at most with few sensilla in one group, palpomere 4 without sensilla; (1) in most species at least female with sensilla in more than one group on palpomere 3 and palpomere 4 usually also with at least sensilla clavata, or, in , numerous sensilla clavata at well developed sensillum coeloconicum. 8. Dorsomedian eye elongation: (0) moderately to well developed; (1) absent or very weak. 9. Temporals: (0) inner verticals present or replaced by frontals, usually more outer verticals; (1) inner verticals absent, outer verticals usually few. 10. Antepronotal lobes: (0) broad, collar like, at most slightly narrowed medially; (1) distinctly narrowed medially. 11. Dorsal antepronotals: (0) absent; (1) present. 12. Humeral pit: (0) inconspicuous; (1) consisting of several smaller areas; (2) conspicuous, oval. 13. Dorsocentrals: (0) erect; (1) decumbent. 14. Dorsocentrals: (0) uniserial anterior; (1) bi–multiserial anterior. 15. Dorsocentrals: (0) uniserial posterior; (1) bi–multiserial posterior. 16. Acrostichals: (0) moderately long to long and strong, (1) short, or absent. 17. Acrostichals: (0) starting in front; (1) starting some distance from antepronotum; (2) in centre of scutum. – Absence scored as (?). 18. Acrostichals: (0) simple or absent; (1) often scalpellate. 19. Prealars: (0) 1–5; (1) 6 or more. 20. Supraalar(s): (0) present; (1) absent. 21. Setae of preepisternum and/or anepisternum: (0) present; (1) absent. 22. Scutellars: (0) uniserial; (1) bi–multiserial. 23. Postnotum: (0) bare; (1) sometimes with setae. – Polymorphies are scored as synapomorphies, i.e. the tendency to have setae is regarded as the synapomorphy. Otherwise the character would be uninformative since only some species of the included genera have setae on the postnotum. To score the polymorphies would be the same as deleting the character. 24. Wing spots: (0) absent; (1) present. 25. Wing membrane: (0) with setae; (1) bare. 26. Wing membrane: (0) not to moderately punctated; (1) coarsely punctated. 27. Anal lobe: (0) relatively well developed, often protruding; (1) weak, reduced or cuneiform wing. 28. Costa: (0) distinctly extended; (1) moderately to not extended.

29. R4+5: (0) ends above or distal to apex of M3+4; (1) ends proximal to apex of M3+4. 30. Clavus: (0) absent; (1) present and ending in distal half of wing; (2) present and ending in proximal half of wing.

31. Cu1: (0) not sinuous; (1) slightly sinuous; (2) strongly sinuous. 32. VR: (0) <1.06; (1) 1.06–1.40; (2) >1.40. 33. Anal veins: (0) An1 extending well beyond cubital fork and An2 conspicuous; (1) anal veins shorter. 34. R veins: (0) setae present on R, R1 and usually R4+5 in both sexes; (1) setae present on R, absent on R1 and often R4+5 of male, at most absent on R1 in female; (2) setae absent on R of male, present in female; (3) setae absent on R and R1 of both sexes, at most 1 apical seta on R4+5. 35. Squama: (0) with setae; (1) bare.

36. Leg ratio of male (LR1): (0) higher than 0.8; (1) 0.5–0.8; (2) lower than 0.5. 37. Trochanter: (0) without keel; (1) at least fore trochanter with keel.

38. Ta 5: (0) not cordiform; (1) cordiform. 39. Tibial spurs: (0) with distinct lateral denticles; (1) denticles indistinct or absent. 40. Inner tibial spur of hind leg: (0) At least 1/2 as long as outer spur; (1) shorter; (2) absent with also second spur of mid leg absent. 41. Hind tibial comb: (0) well developed, occupying full width of tibia; (1) weak or absent. 42. Hind tibial comb: (0) with less than 13 setae; (1) often conspicuous with 13 or more setae of which some about as long as longest spur. 43. Pseudospurs: (0) present; (1) absent. 44. Sensilla chaeticae of tarsi: (0) present; (1) absent. 45. Pulvilli: (0) present and distinct; (1) absent or vestigial, less than 1/2 claw length. 46. Anal point: (0) absent or small and anterior on tergite; (1) represented by hump-like extension of tergite or if absent represented by some stronger median setae; (2) well set off and posterior on tergite. 47. Anal point: (0) not with spatulate microtrichiae-free apex; (1) often with small or large spatulate, microtrichiae-free apex. 48. Anal point: (0) not very broad and rounded to bluntly triangular; (1) often conspicuously broad and rounded to bluntly triangular. 49. Setae on anal point or posterior on tergite IX: (0) conspicuous, bristle-like to lamellate; (1) moderately developed; (2) short, weak or absent. 50. Tergite IX: (0) normal, not very large; (1) large, covering most of hypopygium. 51. Phallapodeme: (0) not pointed and recurved; (1) pointed and recurved. NNFE391015 8/17/04 2:14 PM Page 26

26 H.F. Mendes et al.

Table 3. Continued

52. Superior volsella: (0) present; (1) absent. 53. Superior volsella: (0) not bare and plate-like; (1) bare and plate-like. 54. Gonostylus: (0) simple; (1) double. 55. Heel of gonostylus: (0) absent; (1) present at least in many species. 56. Transverse sternapodeme: (0) convex; (1) straight or concave; (2) absent, sternapodeme triangular. 57. Oral projections of transverse sternapodeme: (0) strongly developed; (1) weak to moderately developed; (2) absent. 58. Crista dorsalis: (0) evident, triangular or rounded preapical; (1) elongate, low; (2) not evident/weak. 59. Megaseta: (0) present; (1) absent. 60. Virga: (0) consisting of tight cluster of at least six spines or two groups of very strong spines; (1) virga not consisting of cluster or groups of spines. 61. Virga: (0) virga when present consisting of cluster or field of spines or few spines with lateral lamellae; (1) consisting of 1–5 short partly fused, sometimes plate-like spines without distinct lateral lamellae. 62. Virga: (0) virga when present consisting of cluster or field of spines or few spines without lateral lamellae; (1) consisting of 1–3 long, median, usually fused spines and distinct lateral lamellae. 63. Gonocoxapodeme: (0) absent, short and straight or evenly curved and ending at base of gonapophysis; (1) continuing basal of vagina or at least past base of gonapophysis. 64. Female tergite IX: (0) undivided; (1) divided by caudal concavity or notch; (2) divided into two setigerous protrusions. 65. Female gonocoxite IX: (0) large, projecting; (1) moderately developed to reduced. 66. Female gonocoxite IX: (0) with several long setae; (1) with less than three long setae; (2) with some long and some short setae; (3) with short setae only. 67. Gonapophysis VIII: (0) undivided; (1) divided with ventrolateral lobe much smaller and more or less brush-like, or with lobes of about same size; (2) divided with dorsomesal lobe smaller and with anterior rounded projection; (3) divided with dorsomesal lobe narrow, often line-like. 68. Apodeme lobe: (0) not apparent; (1) well developed, but not meeting at mid line and with microtrichiae; (2) meeting at mid line and/or with microtrichiae; (3) fully sclerotized. 69. Number of seminal capsules: (0) 3; (1) 2. 70. Seminal capsules: (0) spherical to ovoid, small or of normal size; (1) large, spherical to elongate ovoid. 71. Seminal capsules: (0) at least partly coloured; (1) often completely pale. – Polymorphies are scored as synapomorphies as no genera have all included species with pale capsules and the character otherwise would be uninformative. 72. Opening of spermathecal ducts: (0) separate; (1) common. 73. Spermathecal ducts: (0) not fused; (1) partly fused ducts before common opening. 74. Bulbs of spermathecal ducts: (0) absent; (1) present. 75. Spermathecal ducts: (0) straight; (1) with bend or loop. Pupa 76. Frontal apotome: (0) without warts or tubercles; (1) with warts or tubercles. 77. Frontal setae: (0) present; (1) absent. 78. Thoracic horn: (0) present; (1) absent. 79. Thoracic horn: (0) not rounded to ovoid; (1) often rounded to elongate ovoid. 80. Thoracic horn: (0) not with bulbous base and narrow apical portion; (1) with. – Eukiefferiella is scored as (1) since most species have a thoracic horn with bulbous base and species without thoracic horn apparently have this secondarily reduced. 81. Thorax: (0) mostly smooth to wrinkled; (1) mostly tuberculose or spinulose. 82. Thorax: (0) without tubercle(s) medially or in front; (1) with. – The polymorphy for Cardiocladius s. lat. is scored as a synapomorphy as otherwise the presence of a tubercle will appear as an autapomorphy for Ichthyocladius. 83. Wing sheath: (0) without pearls; (1) with. 84. Dorsocentrals: (0) anterior two and posterior two (three) paired, anterior three grouped, all in row or 2–3 dorsocentrals only; (1) posterior three grouped or all four together. 85. Tergites II–VIII: (0) without posterior spine, or tubercle row(s), but may have caudal hooklets; (1) some with spines or tubercles. 86. Median field of tergite IV: (0) without discrete spine patch(es) or row(s); (1) sometimes with. 87. Tergite I: (0) without posterior spine row(s); (1) with. 88. Sternites II–VII: (0) without posterior spine row(s), but may have caudal hooklets; (1) some with spines or tubercles. 89. Sternites or sternal conjunctives: (0) without caudal hooklets; (1) at least one with. 90. Male sternite VIII: (0) without posterior spine or tubercle row(s); (1) with. 91. Sternite II or II and III: (0) without anterior or median spine group; (1) with. 92. Tergites and sternites: (0) with single or no posterior row of spines; (1) at least some with double to multiple row of spines. 93. Tergite III: (0) without caudal hooklets; (1) with minute caudal hooklets; (2) with conspicuous caudal hooklets, which may be comb- shaped. – absurdus type without thoracic horn and with pearl row, another type with hooklets on T III and PSB on II and III. Characters 93–97 are likely to be interdependent and the presence of hooklets thus is receiving higher weight than other characters. However, several other characters also are likely to be genetically connected and the hooklets certainly are of significance. 94. Tergite IV: (0) without caudal hooklets; (1) with minute caudal hooklets; (2) with conspicuous caudal hooklets, which may be comb-shaped. 95. Tergite V: (0) without caudal hooklets, although rows of conjunctival spinules may be hook-like anteriorly directed; (1) with minute caudal hooklets; (2) with conspicuous caudal hooklets which may be comb-shaped. NNFE391015 8/17/04 2:14 PM Page 27

New Species of Ichthyocladius 27

Table 3. Continued

96. Tergite VI: (0) without caudal hooklets, although rows of conjunctival spinules may be hook-like anteriorly directed; (1) with minute caudal hooklets; (2) with conspicuous caudal hooklets. 97. Tergite VII: (0) without caudal hooklets, although rows of conjunctival spinules may be hook-like anteriorly directed; (1) with minute or conspicuous caudal hooklets. – Polymorphies are scored as synapomorphies as the character otherwise would be uninformative (See character 23). 98. Caudal hooklets of tergite II: (0) present, (1) absent. 99. Pedes spurii B: (0) present; (1) absent. 100. Pedes spurii A on sternite IV: (0) present; (1) absent. 101. Pedes spurii A on sternite VI: (0) present; (1) absent. 102. Tergal conjunctives or terga posterior of tergal spines: (0) without spinules, but may have hooklets in single row; (1) with spinules which may be hooklet-like recurved. 103. Spinules on tergal conjunctives: (0) absent or not hooklet-like recurved or anteriorly directed; (1) hooklet-like recurved or anteriorly directed spinules in about three rows. 104. Taeniate L setae: (0) present; (1) absent. 105. Anal lobe: (0) not extended into projections; (1) extended posteriorly into cylindrical projections with macrosetae at apex. 106. Anal lobe: (0) not with apical projections or extensions; (1) at least sometimes with apical spurs or extended distal of macrosetae. 107. Apical spines of anal lobe: (0) absent; (1) present. 108. Anal lobe: (0) with fringe of setae; (1) without fringe of setae. 109. Inner margin of anal lobe: (0) without long seta; (1) with. 110. Anal macrosetae: (0) subequal; (1) differing; (2) absent or one only. 111. Anal macrosetae: (0) not conspicuously curved and scimitar-like; (1) sometimes conspicuously curved and scimitar-like. 112. Anal macrosetae: (0) not short and spine-like, but may be short and hair-like or absent; (1) short and spine- or thorn-like; (2) absent. 113. Width of anal macrosetae or apical spines: (0) less than 5 mm; (1) 5 mm or more. 114. No. of anal macrosetae: (0) three or more; (1) 2; (2) 0–1. Larva 115. Antenna: (0) with 6–7 segments; (1) with five; (2) with 3–4. 116. Ultimate antennal segment: (0) normal; (1) whip- or thread-like. 117. Antenna: (0) reduced, less than half mandible length; (1) 1/3 as long as head capsule or shorter, but not reduced; (2) longer. 118. Second antennal segment: (0) undivided, fully sclerotised; (1) divided or partly unsclerotised. 119. Lauterborn organs: (0) moderately large to well developed; (1) weak or absent. 120. Antennal blade: (0) short to moderate length, shorter than flagellum when antenna of normal length; (1) conspicuous, longer than flagellum except when flagellum extremely long. 121. S I: (0) plumose, branched, pectinate or palmate; (1) bifid or simple. 122. S I: (0) not bifid; (1) bifid. 123. S I: (0) not simple; (1) simple. 124. Labral lamella: (0) with pectinate, plumose or rugose apex, mostly well developed; (1) weak, no apical teeth or plumosity; (2) absent. 125. Chaetulae laterales: (0) simple or reduced; (1) at least one serrated or plumose. 126. Premandible: (0) simple; (1) with two or more teeth. 127. Premandibular brush: (0) present; (1) absent. 128. Mandible: (0) with 2–3 inner teeth; (1) with four or more. 129. Mola of mandible: (0) smooth; (1) with teeth or spines. 130. Seta interna of mandible: (0) with smooth, slightly plumose laterally or apically, or serrate branches; (1) branches conspicuously branched; (2) seta interna absent. 131. Median tooth of mentum: (0) single; (1) double, bifid or with several median teeth. 132. Mentum: (0) median group of 4–20 teeth not forming straight line on strongly curved mentum; (1) median group of 4–20 teeth forming straight line on strongly curved mentum so that lateral teeth not are visible on uncompressed larvae. 133. Teeth of mentum: (0) 15 or more teeth; (1) 11–14 teeth; (2) fewer. 134. Lateral teeth of mentum: (0) outer tooth not larger or higher than one of the inner teeth; (1) clearly larger or higher. 135. Ventromental plates: (0) well developed, extending past lateral teeth on flattened mentum; (1) reduced or weak. 136. Ventromental plates: (0) without setae (beard) underneath; (1) with setae underneath. 137. Setae submenti: (0) situated at level of base of outer lateral tooth or higher; (1) lower. 138. Claws of anterior parapods: (0) with relatively distinct teeth; (1) smooth or teeth very indistinct. 139. Setal tufts on abdomen: (0) absent; (1) often present. 140. Procercus: (0) well developed; (1) reduced or absent. 141. Anal setae: (0) five or more setae, none conspicuously long; (1) 3–4 setae, none conspicuously long; (2) 0–2 not conspicuously long anal setae; (3) two or more setae with one or two conspicuously long. 142. Supraanal setae: (0) weak, shorter than 1/2 length of anal setae when these well developed or shorter than 2/3 when anal setae short; (1) well developed, longer than 1/2 length of anal setae when anal setae long, longer than 2/3 when anal setae short. 143. Posterior parapods: (0) well developed; (1) small, digitiform; (2) absent to weak but not digitiform. 144. Anal tubules: (0) at least 1/2 length posterior parapods; (1) shorter than 1/2 length posterior parapods or these absent; (2) conspicuously long and narrow. NNFE391015 8/17/04 2:14 PM Page 28

28 H.F. Mendes et al. = 2&3, H = 0&3, G = 001000A 0&2, F = 00010B0B 0100011B0B 1&2&3, E AAAA AAA000100AA = 100100010A0A 1&2, D = 0&1&2, C = AAA0 A 0&1, B = AAA00A1A00AA010001000A AAA00A10001 Orthocladiinae. Polymorphies: A Orthocladiinae. Polymorphies: 012345678901234567890123456 00020AA100AA1000110100A00AA1 000001A0120AA111100010111100110CAC 00010100000001?0111100110100010001 000A0A00000A0110A1AA001AA1000A0A0C AAA000A000001100010A1000 AAA1 0 0 AA0 0 A1 AA1 0 0 AAAA1 A1 A 0000011A000000111001000A01A000111A1 AAA1 0 0 AA0 0 A1 AA1 0 0 AAAA1 A1 A0 A 1&3. = BA0?000111000001?001100010100021111C BA000010000000AA000A1A000A0100110001 200?10001100000010?01000101111020312 123456789111111111122222222223333333 110?10?10000100100011001101000011?01 000000001000E0?A100 000010011100000100011000100100000111 BA01A00AA0000001E001100010A1A001AD01 1100000000000001?001100010011202021A 0010000101000?A110111000011000A1001C A00000001000000120010000A01A000C0001 00010000000000 2100010100000101?0A01100110111110310 200?10?111000001000110011?1100010011 000A000100A00A010?0?1100110100A00AA1 0011000100100A11?0101100010100010001 000A0000A00000 0&2&3, J Fittkau Kieffer B A A 0 A 0 1 A v.d. B v.d. Kieffer Kieffer Sæther A Fries A A Winnertz B v.d. Wulp v.d. = Kieffer Eaton B A Ferrington Ferrington v.d. Wulpv.d. B oi A000100100000111?011011A101000211111 Gowin Sæther & 0 A Kieffer Thienemann Goetghebuer & Sæther Wulp Wiedenbrug & Wiedenbrug Fittkau Halvorsen Spärck Thienemann Thienemanniella Goetghebuer Spärck Boothroyd s. l. Thienemann Physoneura Metriocnemus Table 4.Table Character states for characters 1–144 in some genera of 0&1&2&3, I Character no. Oliveiriella Dratnalia Cricotopus Eukiefferiella Hydrobaenus Ichthyocladius Gen. n. near Heterotanytarsus Kaniwhaniwhanus Corynoneura Chaetocladius Chaetocladius Antillocladius Cardiocladius NNFE391015 8/17/04 2:14 PM Page 29

New Species of Ichthyocladius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ieffer Fittkau Kieffer Kieffer Kieffer Sæther Kieffer A A Fries Winnertz Kieffer Sæther B v.d. Wulpv.d. 0 0 A C oi 001A001A1E0020010000A201010200311001 Gowin Sæther & Kieffer A A Kieffer s. l. Thienemann Halvorsen Thienemann Thienemanniella Goetghebuer Spärck Spärck Boothroyd Goetghebuer Edwards Thienemann & Harnisch Epler & De la Rosa Kieffer Chaetocladius Chaetocladius Corynoneura Cricotopus Dratnalia Propsilocerus Tvetenia Cardiocladius Diplocladius Antillocladius Antillocladius Brillia Bryophaenocladius Eukiefferiella Gen. n. near Gymnometriocnemus Heleniella Heterotanytarsus Heterotrissocladius Hydrobaenus Ichthyocladius Kaniwhaniwhanus Character no. Tempisquitoneura Thienemanniella NNFE391015 8/17/04 2:14 PM Page 30

30 H.F. Mendes et al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ieffer 0 A 1 B Kieffer v.d. v.d. Kieffer Sæther Kieffer Winnertz 0 0 ? v.d. Wulp v.d. Eaton Ferrington Ferrington Sæther v.d. Wulpv.d. 0 A A 0 A A 1 0 A 0 A Continued Kieffer Kieffer Wulp & Wiedenbrug Fittkau & Sæther Thienemann Goetghebuer Goetghebuer Edwards Thienemann & Harnisch Epler & De la Rosa Kieffer s. l. Thienemann Table 4. Table Limnophyes Oliveiriella Orthocladius Propsilocerus Tvetenia Metriocnemus Parakiefferiella Parametriocnemus Physoneura Psectrocladius Pseudosmittia Rheocricotopus Tempisquitoneura Thienemanniella Tokunagaia Character no. Pseudorthocladius Chaetocladius Chaetocladius Corynoneura Cricotopus Antillocladius Antillocladius Brillia Bryophaenocladius Cardiocladius NNFE391015 8/17/04 2:14 PM Page 31

New Species of Ichthyocladius 31 101A1 AAA010001 AAAAAA AAA00E2200A11100A0001 010A00AA100000A00000 000A00A110A000A000000A0A1A00100 0010010101001010122221011100A0000 00100000A100001A10000011111010001 1100000000000000000000000000000000 AAAA 101100001000000010000000000001101001 00001001101010A1010000000111100A0001 0010A1000000100 0010A000A00AA0000000000001111A010001 00000000100010000000000000A000000000 0110000010A000A001000000000000000000 0110A1001001100A0A000000011 000011000100001010002220001110010001 ???000000000000000002220011110000001 ???000001000100000012200001100010001 01A00A1000000000000000000100A1010111 011A1000A01010A101A0000001000001001A ???????????????????????????????????? 0000A000A0000000000000000000AA000000 00101100000010A101A00000011110010011 0A1AA100000100000000000001111AA10001 0010A000000A110A0001000000A001A0000A 00001100A0101001000A1111101110010000 100001000010A0AA10A01111111110000000 001000010000100001010220011110010001 00100001A01010A1010122EE011110010011 Kieffer 0 0 A 1 A Fittkau v.d. v.d. Kieffer Fries 0 A v.d. Wulpv.d. 0 Kieffer Eaton 0 0 A Ferrington Sæther oi 000100000000100101100000011101011011 Gowin Sæther & Kieffer Halvorsen Thienemann Thienemanniella Goetghebuer Spärck Spärck Boothroyd Wulp Wiedenbrug & Wiedenbrug Fittkau Thienemann Goetghebuer & Sæther Goetghebuer Edwards Thienemann & Harnisch Epler & De la Rosa Kieffer Dratnalia Diplocladius Eukiefferiella Gen. n. near Gymnometriocnemus 0 0 A Heleniella Heterotanytarsus Heterotrissocladius Hydrobaenus Ichthyocladius Metriocnemus Kaniwhaniwhanus Limnophyes Oliveiriella Parakiefferiella Orthocladius Parametriocnemus Physoneura Propsilocerus Psectrocladius Pseudosmittia Rheocricotopus Tempisquitoneura Thienemanniella Tokunagaia Tvetenia Pseudorthocladius NNFE391015 8/17/04 2:14 PM Page 32

32 H.F. Mendes et al. 010101A010101100010A AAA00A00A01AA010000A AAA0 AA ?????????????????????????????? 020A010000001AA021100100101A001 010010100A000AA10A01A0C0A01000000A 000BC0A0A01AA2AA10A00010101010000A 000B1010 111111111111111111111111111111111111 011111111112222222222333333333344444 000000101000000011110010100100000001 901234567890123456789012345678901234 00011A1000110002111100A0211011012001 000000201100000011110010111010000001 00000210100A1012011002A02A1011012021 00000010100010120011110010101?01A001 100000202010101211010E1010101?001000 100000101010101201100211101010000000 0A000EC010001012001A10A0C0101A00AA01 12000210C000000200010010111010001001 00000C10100A1012011002102A1011012021 000000202001000111100010100000000A00 00000001C01A0001011A01A0100010000000 000000001000000101100010100010000A00 020202200000110?000A000020111?000000 00000010101A00020100001010101A00010A 000110 00000EC010001102 000000111010101C0A1000A010000?00000A 0000001010000001A1100A10100001000100 ???????????????????????????????????? Fittkau ? A 1 0 A Kieffer 0 A v.d. 0v.d. A Kieffer Kieffer Sæther Fries Winnertz v.d. Wulp v.d. Kieffer Eaton v.d. Wulpv.d. 0 A oi 000000101101000201100110111000000101 Gowin Continued Sæther & Kieffer Thienemann s. l. Halvorsen Thienemann Thienemanniella Goetghebuer Spärck Spärck Boothroyd Wulp Wiedenbrug & Wiedenbrug Fittkau Thienemann Goetghebuer & Sæther Ferrington Table 4. Table Character no. Dratnalia Corynoneura Diplocladius Antillocladius Antillocladius Brillia Bryophaenocladius Cardiocladius Chaetocladius Cricotopus Eukiefferiella Gen. n. near Gymnometriocnemus Heleniella Heterotanytarsus Heterotrissocladius Hydrobaenus Ichthyocladius Kaniwhaniwhanus Limnophyes Metriocnemus Oliveiriella Orthocladius Parakiefferiella Parametriocnemus Physoneura NNFE391015 8/17/04 2:14 PM Page 33

New Species of Ichthyocladius 33

34; 29–46, 35 on S IV–VII as 27–39, 33; 30–41, 34; 29–42, 34; 26–39, 33. Macrosetae as in Fig. 35. Male genital sac extending past anal segment by 38– 83mm (3), female genital sac reaching apex of anal segment.

Phylogenetic analysis A generic-level analysis based on the characters presented in Table 3 and the data matrix in Table 4 with Diplocladius Kieffer as outgroup was performed. In addition to Ichth- yocladius, all genera with caudal hooklets on tergites other than tergite II were included, i.e., genera near Eukiefferiella Thienemann and Corynoneura. Presumed primitive genera such as Brillia Kieffer, Diplocladius Kieffer, Propsilocerus Kieffer, Heterotanytarsus Spärck, and Hydrobaenus Fries were included in order to give direction of trends. These genera also include genera with well developed, finely pecti- nate labral lamellae. Additional common, non-aberrant genera representing different groups of genera also were included. Oliveiriella Wiedenbrug & Fittkau according to the description (Wiedenbrug & Fittkau, 1997) has no supraalar, uniserial scutellars, the shorter hind tibial spur is more than half as long as the outer, and pseudospurs and sensilla chaet- icae are not mentioned. However, M. Spies (personal com- munication) has re-examined the genus and found that a supraalar is present, scutellars are partly bi-serial, the shorter tibial spur is less than half as long as the longer one, pseu- dospurs are absent and sensilla chaeticae present. The new genus near Thienemanniella includes among others Thiene- manniella semifimbriata Sæther (Sæther, 1981: 32) and this type is excluded from Thienemanniella for the characters scored here. The characters are coded ordered to indicate the phylogenetic position of Ichthyocladius relative to known genera. The relationships among more basally placed genera thus should not be taken for relevant. Analyses were done with characters 30, 31, 34, 115, and 124 ordered. When no characters were given weight, nine trees were obtained each with a length of 588 steps, a con- sistency index (CI) of 0.30, a retention index (RI) of 0.46 and a rescaled consistency index (RC) of 0.14 (Fig. 36). When the results were reweighted according to RC, a single tree with 590 steps after resetting to equal weights, CI-0.47, RI-0.70 and RC-0.33 was obtained (Fig. 37). The 100E1010101012A01A001010101000000A 000E101010000200100EA010010100A000 Corynoneura group of genera appears monophyletic and includes Ichthyocladius, while most genera of the Eukief- feriella group are included basally in the same monophyletic 000000C01010000011110010A00000000A00 00000C1010AA0002001001102A100100301E 00000B2000111102110A0E00B0101A01C020 000000101000AA02001000A0100100000A00 020202101011110210010100101010001000 10000010200010121001011010101?00A000 10000010100A0002A01010A0101011000100 group. Dratnalia, Tvetenia and Oliveiriella Wiedenbrug et Fittkau fall outside the group. When the results are reweighted according to the rescaled consistency index Kieffer 0 E Kieffer Tvetenia and Oliveiriella forms a separate group outside the Sæther 0 A

Kieffer main group by two nodes. The Bremer supports are shown above each branch. Several characters obviously are more important than Goetghebuer Edwards Thienemann & Harnisch Epler & De la Rosa Kieffer Propsilocerus Tvetenia Psectrocladius Pseudosmittia Rheocricotopus Tempisquitoneura Thienemanniella Tokunagaia Pseudorthocladius others and should be weighted accordingly. Such weighting NNFE391015 8/17/04 2:14 PM Page 34

34 H.F. Mendes et al.

Figs. 36–39. Parsimony analyses of the relationships between Ichthyocladius Fittkau and some selected Orthocladiinae. (36) Strict consen- sus tree of the 12 trees obtained with no characters weighted. (37) The single tree obtained with characters reweighted according to the rescaled consistency index (RC). (38) Strict consensus tree of the five trees obtained with some characters weighted. (39) The single tree obtained when this result reweighted according to RC.

is rather subjective. However, with experience from related 46, 83, 100, 101, 108, 122, 123, 127 a weight of 5. (Charac- groups and genera and giving weight to characters found ters 93 and 95–97 are not given extra weight since charac- important as synapomorphies in other studies, characters 30, ters 93–97 most likely are not genetically independent and 50, 54, 69, 80, 89, 94, 98, 99, 102, 104, 109, 135 and 136 the presence of caudal hooklets is thus already weighted.) were given a weight of 10; characters 7, 11, 18, 31, 35, 45, The resulting five trees obtained (Fig. 38) had 605–610 steps NNFE391015 8/17/04 2:14 PM Page 35

New Species of Ichthyocladius 35

after resetting to equal weights, CI-0.35, RI-0.61 and RC- Sæther OA (1980): Glossary of chironomid morphology termi- 0.21. The Bremer supports above each branch are number of nology (Diptera: Chironomidae). Entomol Scand Suppl 14: steps higher than 610 after resetting to equal weights. 1–51. Reweighting according to RC resulted in a single tree of 599 Sæther OA (1981): Orthocladiinae (Diptera: Chironomidae) steps, CI-0.45, RI-0.73 and RC-0.36 (Fig. 39). The weighted from the British West Indies, with descriptions of Antillo- result has each of the Corynoneura and Eukiefferiella groups cladius n. gen., Lipurometriocnemus n. gen. Compteros- separately monophyletic before reweighting. The combined mittia n. gen. and Diplosmittia n. gen. Entomol Scand Suppl group is monophyletic after reweighting with the Eukief- 16: 1–46. feriella group basal and paraphyletic. Sæther OA (1986): The myth of objectivity – post-Hennigian The Bremer support is low when the characters are deviations. Cladistics 2: 1–13. unweighted and not reweighted, high for the inclusion of Sæther OA, Kristoffersen L (1996): Chironomids with ‘M-fork’. Ichthyocladius in the Corynoneura group when the charac- A reevaluation of the wing venation of the Corynoneura ters are weighted or reweighted. The phylogenetic placement group (Insecta, Diptera, Chironomidae). Spixiana 19: of Ichthyocladius appears to be confirmed, while the sug- 229–232. gested relationship of the Eukiefferiella and Corynoneura Svensson BS (1986): Eukiefferiella ancyla sp.n. (Diptera: Chi- group is strongly indicated. ronomidae) a commensalistic on Ancylus fluviatilis Müller (Gastropoda: Ancylidae). Entomol Scand 17: 291–298. Wiedenbrug S, Fittkau EJ (1997): Oliveiriella almeidai Acknowledgements (Oliveira, 1946), gen. nov., comb. nov., from South America We are very grateful to Marion Kotrba who provided and with description of the pupae. Spixiana 20: 167–172. facilitated the access to the Zoologische Staatssammlung col- lection of Chironomidae in Munich. The authors would like to thank E.J. Fittkau and M. Spies for refereeing the paper and giving many helpful suggestions. The first author was supported by the Brazilian Research Council of the São Paulo State (FAPESP proc. 00/05903-9 and proc 98/05073- 4 Biota) while completing the paper.

References Cranston PS, Oliver DR, Sæther OA (1983): The larvae of Orthocladiinae (Diptera: Chironomidae) of the Holarctic region – Keys and diagnoses. In: Wiederholm T, ed., Chi- ronomidae of the Holarctic region. Keys and diagnoses. Part 1. Larvae. Entomol Scand Suppl 19: 149–291. Epler JH, De la Rosa CL (1995): Tempisquitoneura, a new genus of Neotropical Orthocladiinae (Diptera: Chironomidae) symphoretic on Corydalus (Megaloptera: Corydalidae). J N Am Benthol Soc 14: 50–60. Fittkau JE (1974): Ichthyocladius n. gen., eine neotropische Gattung der Orthocladiinae (Chironomidae, Diptera) deren Larven epizoisch auf Welsen (Astroblepidae und Loricari- idae) leben. Ent Tidskr Suppl 95: 91–106. Gonser T, Spies M (1997): Southern hemisphere Symbiocladius (Diptera, Chironomidae) and their mayfly hosts (Ephemeroptera, Leptophlebidae). In: Landolt P, Sartori M, eds., Ephemeroptera & Plecoptera: Biology – Ecology – Systematics, MTL – Fribourg, pp. 455–466. Melo AS, Froehlich CG (2001): Macroinvertebrates in neotrop- ical streams: richness patterns along a catchment and assemblage structure between 2 seasons. J N Am Benthol Soc 20: 1–16. Sæther OA (1977): Female genitalia in Chironomidae and other : morphology, phylogenies, keys. Bull Res Board Can 197: 1–211. NNFE391015 8/17/04 2:14 PM Page 36