The Baetodes Complex (Ephemeroptera:Baetidae), Phylogeny, Biogeography, and New Species of Mayobaetis

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The Baetodes complex (Ephemeroptera:Baetidae), phylogeny, biogeography, and new species of Mayobaetis

Article in Freshwater science · December 2015 DOI: 10.1086/684859

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Carolina Nieto1,2

1CONICET-Instituto de Biodiversidad Neotropical, Facultad de Ciencias Naturales and Instituto Miguel Lillo, Miguel Lillo 251, Tucumán, Argentina, UNASUR (4000)

Abstract: The Baetodes complex was erected by Lugo-Ortiz and McCaﬀerty based on 2 characters: tarsal claws in the larva with a strong subapical seta and forceps in the male adult with basal segments with a distomedial projection. This complex in- cludes the genera Baetodes, Lugoiops, Mayobaetis, Moribaetis, Prebaetodes,andSpiritiops. It is restricted to the New World, and larvae always live in well-oxygenated streams. A cladistic analysis was conducted for the complex on the basis of a matrix of 16 taxa and 92 morphological characters, all derived from external morphology of adults and larvae. A biogeographic analysis was carried out through Spatial Analysis of Vicariance to establish possible vicariant events that generated this taxonomic complex. The cladistic analysis under implied weights yielded 3 shortest trees, and the Baetodes complex was recovered as monophyletic. The biogeographic analysis reconstructed 3 disjunction nodes when no nodes were removed and 2 more when 1 node was removed. The vicariant events cannot be attributed unquestionably to any paleoclimatic or microtectonic event, but they coincide under certain scenarios. Mayobaetis yala, new species, from Argentina, was described. It can be distinguished from other species in the genus because segment I of the forceps lacks a projection in the male imago and the dorsal surface of the abdomen lacks a medial dark line in the larva. Key words: Baetodes complex, systematic, biogeography, Neotropical region

The Baetodes (Ephemeroptera:Baetidae) complex, a group distributed from Arizona and Texas to Argentina (Koss restricted to the New World, was erected by Lugo-Ortiz 1972, McCafferty and Provonsha 1993, Nieto 2004, de Souza and McCafferty (1996) to include 4 genera: Baetodes Need- et al. 2011). Lugoiops is a monotypic genus from Guatemala, ham and Murphy (1924), Mayobaetis Waltz and McCafferty whereas Moribaetis has 3 well-known species from Cen- (1985), Moribaetis Waltz and McCafferty (1985), and Pre- tral America and Mexico (Waltz and McCafferty 1985). baetodes Lugo-Ortiz and McCafferty (1996). The proposed Moribaetis mimbresaurus, a species from the USA was de- complex was based on 2 characters: tarsal claws in the larva scribed by McCafferty (2007) on the basis of adults, so it with a strong subapical seta and forceps of the male adult was not included in the analysis. Three species from South with basal segments with a distomedial projection (Lugo- America were transferred to this genus by Lugo-Ortiz and Ortiz and McCafferty 1996). Later, McCafferty and Baum- McCafferty (1999), but these species were later transferred gardner (2003) described Lugoiops and placed it within to another genus or considered nomen dubium (Cruz and the complex. In 2010, Nieto proposed a cladistic hypothe- Salles 2014). Prebaetodes has 2 known species from Colom- sis of the family Baetidae in South America, and the genus bia, Ecuador, and Venezuela (Chacón et al. 2010). Spiritiops Spiritiops was recovered as a member of the complex. silvudus Lugo-Ortiz and McCafferty (1998) was the only spe- The evolutionary radiation responsible for the Baetodes cies in this widely distributed genus in Brazil, French Guiana, complex occurred in the Neotropical region (McCafferty Surinam, and Venezuela, until Nieto and Derka (2012) de- 1998). McCafferty (1998) suggested that this complex orig- scribed a 2nd species from the Tepui region of Venezuela. inated in South America. Several genera were collected in Mayobaetis was originally placed by Waltz and McCafferty waterfalls (Mayo 1973), which may represent the ancestral (1985) as a subgenus of Moribaetis that included 1 spe- or generalized habitat of the group. The larvae have been cies, Baetis ellenae (Mayo 1973). However, Lugo-Ortiz and collected on the surface of rocks in rapids or in the splash McCafferty (1996) assigned generic rank to Mayobaetis. zones (Nieto and Derka 2012) and in fast-flowing water, Since then, this genus has been recorded from Argentina, always in well-oxygenated streams (Salles et al. 2004). Bolivia, Colombia, Costa Rica, Ecuador, and Peru. The genus Baetodes isoneofthemostdiverseandwidely The present biogeographic analysis is an attempt to rec- distributed genera in the family and has 44 known species ognize the barriers separating a biota rather than to propose

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DOI: 10.1086/684859. Received 24 July 2014; Accepted 21 September 2015; Published online 15 December 2015. Freshwater Science. 2016. 35(1):000–000. © 2016 by The Society for Freshwater Science. 000 000 | The Baetodes complex C. Nieto relationships between predefined areas. The method used paper. As in a recent analysis by Nieto (2010), 5 species here follows the approach advocated by Hovenkamp (1997, were included as outgroups. Four of them belong to the 2001) because the analysis is focused on taxon history (Arias family Baetidae: Rivudiva minantenna, Rivudiva tricho- et al. 2011). Given a phylogeny, taxon history methods are basis, Cryptonympha copiosa, and Cryptonympha dasilvai. used to explain the distribution of a particular taxon in a The trees were rooted with Metamonius anceps (Nesame- geological context. In other words, a Spatial Analysis of Vi- letidae), considered closely related to Baetidae (e.g., Staniczek cariance (SAV; Arias 2010) is an attempt to discover disjunct 1997, Kluge 2004). A detailed list of specimens of these spe- (allopatric or vicariant) distribution among sister groups. cies can be found in Nieto (2010). Other species included Thus, knowing the phylogeny of the taxa is crucial. were S. tepuensis, scored from type material (IML), and The main goals of the present analysis are to: 1) ex- P. meridinensis scored from the literature (Chacón et al. plore the phylogenetic relationships among the genera of 2010). the Baetodes complex, based on a previous cladistic analysis (Nieto 2010) and the new species described since then; 2) perform a Spatial Analysis of Vicariance to establish the Characters possible vicariant events that affected this complex; and All characters in this study (Appendix S2) were de- 3) describe a new species of Mayobaetis, based on larvae rived from the external morphology of adults and larvae, and adults from Argentina. and most of them were adapted from Nieto (2010). Eighty characters were included in the analysis. Most were derived from larval morphology (62) and the rest (18) from the MATERIAL AND METHODS male imago. Fifty-five characters were binary and 25 were Larvae were collected with kick and D-frame nets. Light multistate. All characters were treated as nonadditive (un- ffi traps were most e cient for capturing adults, but rearing ordered). larvae provided an accurate association of both states. All specimens were conserved in 96% ethyl alcohol. To study Phylogenetic analysis morphology, mature larvae and imagos were selected and dissected. Dissected parts of the specimens under study The matrix was analyzed with the program TNT (Golo- boff et al. 2008) under an implied weights assumption (Golo- were mounted on microscope slides with Canada balsam. ff k Line drawings were made with a camera lucida attached to bo 1993) with values ranging from 2 to 6. The shortest a microscope. Photographs of adult and larval specimens trees were obtained under Analyze/TraditionalSearch com- mands; i.e., 500 replicates of random addition sequences were taken with an SMZ-10 stereomicroscope (Nikon, Tokyo, – Japan or a BX-51 microscope; Olympus, Tokyo, Japan), with followed by tree bisection reconnection, saving 10 trees/ a Nikon D5000 digital camera. For some of the pictures, a replication. Group support was obtained by applying frequency differences (Goloboff et al. 2003), which were cal- series of partially focused images was processed with the fi program Combine ZP (Combine Z5; Hadley 2009) to pro- culated with 500 jackkni ng replicates (Farris et al. 1996) duce final images with enhanced quality. The material ex- by symmetrical resampling of the original matrix (each character presents the same probability of being positively or amined is housed at the following institutions: IBN (Instituto ff de Biodiversidad Neotropical), Tucumán, Argentina; IML negatively reweighted; Golobo et al. 2003). Each jackknif- (Instituto Miguel Lillo), Tucumán, Argentina; ULUMSA ing replicate was calculated by 10 random-addition sequences plus tree bisection–reconnection, saving 10 trees/replication. (Instituto de Ecología, Unidad de Limnología, Universidad Mayor de San Andrés), La Paz, Bolivia; MEUV (Museo de Entomología de la Universidad del Valle), Cali, Colombia. Biogeographic analysis All available geographical records of the Baetodes com- Taxa plex were mapped with DIVA-GIS (Hijmans et al. 2012) A matrix of 16 taxa and 80 morphological characters based on the limits of the freshwater ecoregions of the was constructed (Appendix S1). The ingroup taxa belong- world by Abell et al. (2008). Water lines and water bodies ing to the Baetodes complex that were included in the were provided on the DIVA website (http://www.diva-gis matrix are: Lugoiops (L. maya), Prebaetodes (P. sitesi and .org/Data). The biogeographic analysis was carried out through P. meridinensis), Spiritiops (S. silvudus and S. tepuensis), SAV (Arias et al. 2011), a method that uses observed dis- Moribaetis (M. macaferti, M. maculipennis, and M. sal- tributions as data. SAV does not assume predefined areas vini), and Mayobaetis (M. ellenae and M. yala, new spe- or a hierarchical relationship between them. The main pur- cies). Moribaetis mimbresaurus was not included because pose of this method is to discover disjunct (allopatric or only the adults are known and lack of information on larvae vicariant) distributions among sister groups. The analysis would have left several gaps in the matrix. Baetodes was was implemented in the Vicariance Inference Program (VIP; included at the generic level because it has >40 species, Arias 2010). Distributions were represented as absence/ and their inclusion would have exceeded the scope of this presence data in 1° × 1° cells in a latitudinal–longitudinal grid Volume 35 March 2016 | 000

(maximum ﬁll = 0) on a National Aeronautics and Space Agency (NASA) World Vegetation map (Stöckli 2015; NASA’s Earth Observatory Group). A heuristic search (it- erations: n = 1000) was conducted under the default set- tings to obtain the best reconstructions of disjunct pairs. Searches using diﬀerent grid sizes (2° × 2°) and costs were conducted for comparative purposes, but the results were similar.

RESULTS AND DISCUSSION Phylogeny The analysis under implied weights yielded 3 shortest trees (length = 168 steps, fit = 53.45). Figure 1 shows the strict consensus of the shortest trees with Metamonius anceps astheoutgroup.Thetopologyoftheconsensustree is similar to that in fig. 73 by Nieto (2010). Figure 2. Vicariant events obtained from the Vicariance In- The Baetodes complex was obtained as monophyletic ference Program (VIP) analysis. Black squares indicate disjunct sister pairs. group (Node A; Fig. 1, Appendix S3). Four synapomorphies in the larvae and good support values under jackknifing sus- tained this result: the glossae overlapped to paraglossae (char- was recovered with 8 synapomorphies, among which pedicel acter 29), tibiae longer than tarsi (character 43), tarsal claws of the antennae with scale-like tuberculations (character 3), with strong subapical setae (character 48), and abdominal dorsoventrally flattened scape (character 4), long scape (>3× terga with covers (character 50). The Baetodes complex the pedicel) (character 6), and cerci (Fig. 6L) with spines on originally did not include Spiritiops. Nieto (2010) showed external margins (character 60), were particularly important. that this genus must be included in the complex to sup- Prebaetodes (Node D; Fig. 1, Appendix S3) displayed 6 syn- port its monophyly. This result was confirmed in this analy- apomorphies, such as lingua without projection (charac- sis. The strong subapical seta in the tarsal claws, which is ter 21), long and slender teeth at the apex of the maxilla the main synapomorphy for this group, is present in other crown (character 27), and absence of gill tracheation (char- genera, such as Echinobaetis, Indobaetis,andPapuanatula, acter 55). Spiritiops (Node B; Fig. 1, Appendix S3) was re- outside the New World. However, a preliminary analysis covered with 4 synapomorphies: crenelated lateral margin of the family Baetidae suggests that these genera are not of superlinguae (character 22), segment III of the labial palp related to this complex (J.-L. Gattolliat, Musée cantonal (character 35) showing length subequal to width, length of de zoologie, Lausanne, and CN, unpublished data). gill/length of tergum ratio 2.0 to 2.9 (character 52) and fore- The inclusion of Mayobaetis yala, new species, P. wings with pterostigma 3–5 cross-veins (character 69). meridinensis, and S. tepuensis, 3 genera previously consid- Baetodes was recovered as a sister group of Lugoiops ered monospecific, confirmed the monophyly of the Bae- (Node E; Fig. 1, Appendix S3). In turn, Mayobaetis was todes complex. Mayobaetis (NodeF;Fig.1,AppendixS3) recovered as a sister group of both Baetodes and Lugoiops (Node G). Prebaetodes was placed as sister to all the former (Node H), followed by Moribaetis (Node I), and last, Spir- itiops, as a sister group of the rest of the complex. Some characters showed a tendency to diminish in size or number inside the complex. For instance, the basal state in the tarsal claw (character 47) in Spiritiops displays 2 rows of denticles, a feature that is reduced to 1 in the rest of the complex. The antennae (character 5) are fairly long in Spir- itiops, Moribaetis,andPrebaetodes, whereas they are short in Mayobaetis, Lugoiops, and Baetodes. A similar pattern can be observed in the length of the terminal filament (character 59).

Biogeographic analysis The complex is distributed throughout North, Central, Figure 1. Strict consensus of the 3 shortest trees obtained. and South America. It appears mainly in the freshwaters of The numbers above the nodes indicate frequency diﬀerences. the ecoregions Papaloapan, Chiapas, San Juan, and Chiriquí 000 | The Baetodes complex C. Nieto

(Abell et al. 2008) in North and Central America. These as Mayobaetis or Prebaetodes, are distributed mainly in ecoregions are characterized by tropical and subcoastal the west side in the Amazonas High Andes, Amazonas rivers with tropical climate featuring a dry season fol- Lowlands, and in the North Andean and Central Paciﬁc lowed by a rainy season (Abell et al. 2008). In South Amer- slopes. These freshwater ecoregions are composed of ica, the complex is distributed in 2 areas. Some genera, such many independent short rivers in small watersheds. These

Figure 3. Hypothetical barriers generated by the Vicariance Inference Program (VIP). A.—Vicariant node 1, showing the split of Spiritiops (red dots) and the other genera of the complex (blue dots). B.—Vicariant node 2, showing the split of Moribaetis (red dots) and the other genera of the complex (blue dots). C.—Vicariant node 3, showing the split of Prebaetodes meridinensis (red dots) and Prebaetodes sitesi (blue dots). D.—Vicariantnode4,showingthesplitofMayobaetis yala (red dots) and Mayobaetis ellenae (blue dots). E.—Vicariant node 5, showing the split of Baetodes (red dots) and Lugoiops maya (blue dots). Open squares, species not considered in this node. Volume 35 March 2016 | 000 rivers are narrow and high-gradient with a climate that from 1 reconstruction in which the node (Baetodes + L. varies from cool to cold and temperatures inversely pro- maya)(M. yala + M. ellenae) was removed. portional to increasing altitude. In eastern South Amer- A brief description of tectonic background is necessary ica, some genera, such as Baetodes or Spiritiops, are dis- to explain the possible coincidence between vicariant events tributed in the Upper and Lower Paraná, lower Uruguay, of this complex and tectonic events. During the middle and the Amazonas Guiana Shield and Orinoco Guiana Shield, late Miocene, 3 successive Atlantic marine transgressions, and Maracaibo, among other freshwater ecoregions. The all informally known as the Paranaean Sea, were recorded climate is humid subtropical with warm summers. The hab- in South America. The northwestern part of the Paranaean itats range from large low-gradient rivers to steep head- Sea was connected with the so-called “Tethys Waterspout”. water streams with fast waters. During this time, an open seaway separated terrestrial envi- In the biogeographic analysis, without removing nodes ronments or, in this case, freshwater ecosystems. During (Fig. 2), VIP recovered 1 reconstruction with a cost of 9.00 the middle Miocene, a great increase of thermal gradient and 3 disjunction pairs (nodes 3–5). However, Baetodes was occurred from the equator toward both poles. In addition, removed in most of the 175 VIP possible reconstructions the uplifting Andes constituted a major barrier to moisture- in which some nodes were removed, as expected. This out- laden South Paciﬁc winds (Ortiz-Jaureguizar and Cladera come creates a challenge in attempts to ﬁnd a biogeograph- 2006). All of these elements could have concurred with ical explanation because this genus is widely distributed. the split between Spiritiops and the rest of the complex Two more disjunct pairs were recovered (nodes 1 and 2) (Fig. 3A), i.e., node 1. Concurrently, the absence of the

Figure 4. Mayobaetis yala sp. nov., dorsal habitus. A.—Male imago. B.—Female imago. C.—Larva. Pictures processed with the program Combine ZP. 000 | The Baetodes complex C. Nieto

Isthmus of Panama, which was finally closed during the late Pliocene, coincided with the split of Moribaetis from the rest of the genera (Fig. 3B), i.e., node 2. Unlike most of the South American Andes, which were formed by convergence across a conventional type-B sub- duction, the Merida Andes (MA) have some special traits that are worth considering. The MA do not have a direct historical relationship with the rest of the range (Aude- mard 2003). In addition, together with the eastern mountain ranges, they are separated by the southern termina- tion of the left-lateral strike-slip Santa Marta–Bucaramanga fault and the northwest–southeast trending Santander Mas- sif (Audemard 2003). This striking difference would explain the vicariant event that P. meridinensis and P. sitesi (node 3; Fig. 3C) do not share the same geographical distribution even though both inhabit Andean regions at high altitude. Prebaetodes meridinensis occurs in the MA, whereas P. sitesi occurs in the Ecuadorian and Colombian Andes. Another interesting vicariant event took place probably as the result of combined action of the Andean uplift and Figure 5. Mayobaetis yala sp. nov. Male imago. A.—Forewing. the western Antarctic glaciation during the middle Miocene. B.—Hindwing (a) and hindwing detail (b). C.—Genitalia (ventral These 2 events caused an increase in the thermal gradient view). from the equator to the poles (Ortiz-Jaureguizar and Cladera 2006). As a consequence, 2 species, M. yala and M. ellenae dark line (Fig. 4C), 2) lingua narrow and pointed apically (node4;Fig.3D),mayhaveappearedoneithersideofthe (Fig. 6E), 3) body length, short, <8 mm. mountain. A similar result was obtained for the genus Cyriocosmus (Araneae:Theraphosidae) (fig. 11 by Ferretti Male imago (Fig. 4A) Length, body: 8.8–9.0 mm; fore et al. 2012). wings: 9.4–9.5 mm; hind wings: 1.4 mm. Head yellowish The last vicariant event (node 5; Fig. 3E) to be consid- brown, antennae orange brown. Compound eye oval and ered was caused by the extreme geological complexity of orange. Height of stalk subequal to eye diameter, ocelli black. Central America where, in effect, tectonic and volcanic Thorax yellowish brown. Mesoscutelum orange brown, meta- processes operate in tandem (Mann et al. 2007). Central scutelum brownish. Pleurae dark brownish. Pro, meso, and America encompasses an intricate mosaic of dynamic land- metasterna orange brown. Legs yellowish. Leg I: tibia 1.5× scapes shaped by a wide range of Earth surface processes, the length of femur, tarsus subequal to tibia. Legs II and and the resulting physiography of each landscape features III: tibiae 1.5× the length of femora, tarsi ½ the length of a distinctive landform assemblage that preserves a unique femora. Wings hyaline, except costal and subcostal space of history of landscape evolution (Marshall 2007). forewings opaque. Marginal intercalary veins longer than the space between longitudinal veins (Fig. 5A). Hind wings with 2 longitudinal veins (Figs. 5Ba, Bb). Abdomen orange TAXONOMY brown, posterior and lateral margin of terga brownish. Sterna Mayobaetis Waltz and McCafferty yellowish. Genitalia orange brown, forceps 3-segmented, Moribaetis (Mayobaetis) Waltz and McCafferty 1985: segment I without medial projection, segment III short 240. (Fig. 5C). Cerci orange brown. Mayobaetis;Lugo-OrtizandMcCafferty 1996:369; Do- mínguez et al. 2006:161. Female imago (Fig. 4B) Length, body: 9.7–9.8 mm; fore (Type-species: Baetis ellenae Mayo, original designation) wings: 10.6–10.7 mm; hind wings: 1.2–1.3 mm. Head reddish brown. Compound eyes grayish. Thorax reddish Mayobaetis yala Nieto, sp. nov. (Figs 4A–C, 5A–C, 6A–L) brown. Mesoscutelum orange brown. Pro-, meso-, and metasterna reddish brown. Legs yellowish brown. Abdomen Diagnostic characters Mayobaetis yala sp. nov. can be reddish brown. distinguished from the other species of the genus by the following combination of characters. In the male imago: Larva (Fig. 4C) Length, body: 7.7–8.0 mm; cerci: 6.3 mm; 1) segment I of the forceps without projection (Fig. 5C). terminal filament: 1.9 mm. Antennae: 1.8–1.9 mm. Gen- In the larva: 1) dorsal surface of abdomen without a medial eral coloration yellowish brown. Head yellowish brown, Figure 6. Mayobaetis yala sp. nov., larva. A.—Antennae. B.—Labrum dorsal (left) and ventral (right) view (a) and anterior margin (b). C.—Left mandible. D.—Right mandible. E.—Hypopharynx. F.—Maxilla. G.—Labium ventral (left) and dorsal (left) view. H.—Leg I. I.—Tarsal claw. J.—Posterior margin of tergum IV. K.—Gill IV. L.—Cerci. 000 | The Baetodes complex C. Nieto compound eyes of male brownish. Antennae (Fig. 6A) pale curved, a character that was omitted in the original de- yellow, pedicel of the antennae with scale-like tubercula- scription. tions. Mouthparts yellowish brown. Labrum (Fig. 6Ba) wider than long, dorsal surface with a row of subapical setae, ven- Distribution Bolivia, Colombia, Costa Rica, Ecuador, Perú, – trally lateral margin sclerotized and with a row of 3 4 short and Venezuela. spine-like setae, apical setae bifid (Fig. 6Bb). Mandibles (Fig. 6C, D) with incisors fused apically, without setae be- Material examined 15 larvae, BOLIVIA, Depto. La Paz, tween prostheca and mola. Left mandible (Fig. 6C) with San Pedro abajo, cascada camino a Coroico, S 16°15′12′′, prostheca robust and with denticles apically. Right pros- W 67°47′08′′, 2250 m, 25.XI.2000, Domínguez, Molineri theca bifid (Fig. 6D). Hypopharynx with lingua longer than y Nieto cols. 30 larvae, San Pedro arriba, cascada camino superlinguae (Fig. 6E). Maxillae (Fig. 6F) with palpi subequal a Coroico, S 16°16′15′′, W 67°47′05′′, 2400 m, same date in length to galea-lacinia, with 2 segments, segment II with and collectors. 4 larvae, COLOMBIA, Valle del Cauca, a distal constriction. Labium (Fig. 6G) with glossae sub- Municipio de Riofrío, río La Cristalina, 2 km al Occi- equal in length to paraglossae, truncate apically and with dente de Salónica, Cuenca alta del río Riofrío, N 04°07′ 2–3 pectinated setae. Paraglossae with 3 rows of pectinated 39′′, W 76°24′13′v, 1240 m, 26.VIII.2000, Zuñiga, Gri- setae. Labial palpi with segment II without medial projec- sales, R. J. Cardoso-Zuñiga cols, 2 ♂ imagos same local- tion, segment III oval and with short spines. ity: 17.IX.2001, Zuñiga, Echeverri, Cardoso-Zuñiga. One Thorax yellowish brown. Anterior margin of metathorax larva, Municipio de Cali, Corregimiento de Pance, Ver- brownish. Pleurae brownish. Sterna pale yellow. Legs (Fig. 6H) eda El Pato, Cuenca alta Río Pance, 1600 m, N 3°19′52′′, yellowish brown, femora with a pale longitudinal band on W76°38′48′′, 2.VI.2007, Zuñiga, Cardona, López, Mon- dorsal surface, tarsi brownish apically. Tarsal claws (Fig. 6I) taña, col. 12 larvae from Bolivia housed at IBN, the rest of with 8–9 denticles, and with strong subapical setae. the material housed at ULUMSA. 2 larvae and 1 ♂ imago Abdomen orange brown (Fig. 4C), posterior margin of from Colombia housed at IBN, the other material from terga smooth and brownish (Fig. 6J). Sterna yellowish brown. the same country housed at MEUV. Gills (Fig. 6K) whitish, main trachea weak pigmented. Caudal filaments yellowish. Cerci (Fig. 6L) with spines on external margins. CONCLUSION The hypothesis that Spiritiops belongs to the Baetodes complex is reinforced. Three previously monospecific Etymology Yala is the name of one of the rivers where genera, Mayobaetis, Prebaetodes and Spiritiops, were re- this species was collected. covered each as monophyletic. The biogeographic analysis reconstructed 3 disjunction nodes without removing Distribution Argentina, Jujuy. nodes, and 2 more if 1 node is removed. Although some vicariant events and corresponding geological phenomena Material examined Holotype female larva: ARGENTINA, cannot be proved to have taken place at the same time, Jujuy, Yala, cascadita camino a Reyes, S 24°07′25′′, W 65° some striking coincidences may suggest that in effect they 29′18′′, 26.III.1999, Molineri col. Paratypes, 24 larvae, 6 ♂ took place simultaneously. and 9 ♀ imagos, (2 ♂ and 3 ♀ reared) same locality and collector. 5 larvae: Estancia Jakulica, Arroyo El Nogalar, 19. XI.1985, Fidalgo col. Material housed at IBN. ACKNOWLEDGEMENTS The author is indebted to Eduardo Domínguez, Carlos Molineri, and María del Carmen Zuñiga for the loan of mate- Mayobaetis ellenae Mayo rial. Carlos Molineri provided helpful comments and discussions Baetis sp. 1 Roback 1966:137. during this study. The valuable suggestions of Luisa Montivero, Baetis ellenae Mayo 1973:285; Berner 1980:190. Atilano Contreras-Ramos, and an anonymous referee are ap- Moribaetis (Mayobaetis) ellenae; Waltz and McCafferty preciated. The author thanks the Willi Hennig Society for mak- 1985:240. ing the program TNT free and available, and Salvador Arias for Mayobaetis ellenae;Lugo-OrtizandMcCafferty 1996:369; help with VIP. Financial support from Consejo Nacional de Cruz et al. 2012:2. Investigaciones Científicas y Técnicas (CONICET), to which the author belongs, is acknowledged. This research was partially supported by the following grants: PIP 1484 and PIP 0330. Remarks This species was originally described as Baetis fi ellenae (Mayo 1973). Its classi cation has shifted between LITERATURE CITED ff di erent statuses, in a subgenus of Moribaetis or in its own Abell, R., M. L. Thieme, C. Revenga, M. Bryer, M. Kottelat, N. genus, Mayobaetis, but it has never been described again. Bogutskaya, B. Coad, N. Mandrak, S. Contreras Balderas, W. New material assignable to this species was reviewed, and Bussing, M. L. J. Stiassny, P. Skelton, G. R. Allen, P. Unmack, inthecaseofadults,thedistinctiveCuPveinsarestrongly A. Naseka, R. Ng, N. Sindorf, J. Robertson, E. Armijo, J. V. Volume 35 March 2016 | 000

Higgins, T. J. Heibel, E. Wikramanayake, D. Olson, H. L. Hovenkamp, P. 2001. A direct method for the analysis of vicari- López, R. E. Reis, J. G. Lundberg, M. H. Sabaj Pérez, and ance patterns. Cladistics 17:260–265. P. Petry. 2008. Freshwater ecoregions of the world: a new Kluge, N. J. 1997. Classification and phylogeny of the Baetidae map of biogeographic units for freshwater biodiversity con- (Ephemeroptera) with description of the new species from servation. BioScience 58:403–414. the Upper Cretaceous resins of Taimyr. Page 527–535 in P. Arias, J. S. 2010. VIP: Vicariance Inference Program. (Available Landolt and M. Sartori (editors). Ephemeroptera and Plecop- from: http://www.lillo.org.ar/phylogeny/VIP) tera. Biology–Ecology–Systematics. Mauron + Tinguely and Arias, J. S., C. A. Szumik, and P. A. Goloboff. 2011. Spatial analy- Lachat, SA, Fribourg, Switzerland. sis of vicariance: a method for using direct geographical in- Kluge, N. J. 2004. The phylogenetic system of Ephemeroptera. formation in historical biogeography. Cladistics 27:1–12. 442pp. Kluwer, Dordrecht, The Netherlands. Audemard, M. 2003. Geomorphic and geologic evidence of ongo- Koss, R. W. 1972. Baetodes: new species and new records for ing uplift and deformation in the Mérida Andes, Venezuela. North America (Ephemeroptera: Baetidae). Entomological News Quaternary International 101/102:43–65. 83:93–102. Berner, L. 1980. II. Ephemeroptera nymphs. Proceedings of the Lugo-Ortiz, C. R., and W. P. McCafferty. 1996. Phylogeny and Academy of Natural Sciences of Philadelphia 132:187–194. classification of the Baetodes complex (Ephemeroptera:Baetidae), Chacón, M. M., M. L. Pescador, and S. Segnini. 2010. The adult with description of a new genus. Journal of the North Ameri- and redefinition of the genus Prebaetodes Lugo-Ortiz & McCaf- can Benthological Society 15:367–380. ferty (Ephemeroptera: Baetidae), with description of a new Lugo-Ortiz, C. R., and W. P. McCafferty. 1998. Five new genera of species from Venezuela. Aquatic Insects 32(2):143–157. Baetidae (Insecta: Ephemeroptera) from South America. Anna- Cruz, P. V., R. Boldrini, C. Quinto, and H. Frontado. 2012. New les de Limnologie – International Journal of Limnology 34:57–73. Baetidae (Insecta: Ephemeroptera) records from Venezuela Lugo-Ortiz, C. R., and W. P. McCafferty. 1999. Revision of and nymph description of an unnamed Fallceon species. In- South American species of Baetidae (Ephemeroptera) previ- ternational Journal of Zoology 2012. doi:101155/2012/837092 ously placed in Baetis Leach and Pseudocloeon Klapalek. An- Cruz,P.V.,andF.F.Salles.2014.FurthernotesonSouthAmer- nales de Limnologie – International Journal of Limnology 35 ican species of Baetidae (Ephemeroptera) assigned to Moribaetis (4):257–262. Waltz & McCafferty, 1985. Zootaxa 3793(3):398–400. Mann, P., R. D. Rogers, and L. Gahagan. 2007. Overview of de Souza, M. R., F. F. Salles, and J. L. Nessimian. 2011. Three plate tectonic history and its unresolved tectonic problems. new species of Baetodes Needham and Murphy (Ephemeroptera: Pages 205–241 in J. Bundschuh and G. Alvarado (editors). Baetidae) from Espirito Santo State, Brazil. Aquatic Insects 33 Central America: geology, resources, and hazards. Taylor (2):93–104. and Francis, London, UK. Domínguez,E.,C.Molineri,M.L.Pescador,M.D.Hubbard,andC. Marshall, J. S. 2007. Geomorphology and physiographic prov- Nieto. 2006. Ephemeroptera of South America, Aquatic Biodi- inces of Central America. Pages 75–122 in J. Bundschuh and versity of Latin America. Volume 2. Pensoft Press, Sofia, Poland. G. Alvarado (editors). Central America:geology,resources,and Farris, J. S., V. A. Albert, M. Källersjö, D. Lipscomb, and A. G. hazards. Taylor and Francis, London, UK. Kluge. 1996. Parsimony jackknifing outperforms neighbor- Mayo, V. K. 1973. A new species of Baetis from Ecuador (Ephe- joining. Cladistics 12:99–124. meroptera: Baetidae). Pan-Pacific Entomologist 49:285–288. Ferretti, N., A. González, and F. Pérez-Miles. 2012. Historical McCafferty, W. P. 1998. Ephemeroptera and the great American biogeography of the genus Cyriocosmus (Araneae: Therapho- interchange. Journal of the North American Benthological So- sidae) in the Neotropics according to an event-based method and ciety 17:1–20. spatial analysis of vicariance. Zoological Studies 51:526–535. McCafferty, W. P. 2007. Moribaetis mimbresaurus, new species Flowers, R. W. 1987. The adult stage of three Central American (Ephemeroptera: Baetidae): first representative of the genus Baetodes (Ephemeroptera: Baetidae) with notes on the genus. north of Mexico. Proceedings of the Entomological Society Aquatic Insects 9:1–10. of Washington 109:696–699. Gillies, M. T. 2001. A new species of Aturbina (Ephemeroptera, McCafferty, W. P., and D. E. Baumgardner. 2003. Lugoiops Baetidae) Lugo-Ortiz and McCafferty from Uruguay. Page 331– maya, a new genus and species of Ephemeroptera (Baetidae) 335 in E. Domínguez (editor). Trends in Research in Ephem- from Central America. Proceedings of the Entomological So- eroptera and Plecoptera. Kluwer Academic/Plenum Publishers, ciety of Washington 105:397–406. New York, USA. McCafferty, W. P., and A. V. Provonsha. 1993. New species, sub- Goloboff, P. 1993. Estimating character weights during tree search. species, and stage descriptions of Texas Baetidae (Ephem- Cladistics 9:83–91. eroptera). Proceedings of the Entomological Society of Wash- Goloboff, P. A., J. S. Farris, M. Källersjö, B. Oxelman, M. J. ington 95:59–69. Ramírez, and C. Szumick. 2003. Improvements to resampling Needham, J. G., and H. E. Murphy. 1924. Neotropical mayflies. measures of group support. Cladistics 19:324–332. Bulletin of the Lloyd Library of Botany, Pharmacy and Materia Goloboff, P. A., J. S. Farris, and K. Nixon. 2008. TNT, a free pro- Medica, Entomological Series 24:1–79. gram for phylogenetic analysis. Cladistics 24:774–786. Nieto, C. 2004. The genus Baetodes (Ephemeroptera: Baetidae) Hadley, A. 2009. Combine ZP, image stacking software. (Avail- in South America with the description of new species from able from: http://www.hadleyweb.pwp.blueyonder.co.uk) Argentina, Bolivia and Peru. Studies on Neotropical Fauna Hijmans, R. J., L. Guarino, and P. Mathur. 2012. DIVA-GIS, and Environment 39:63–79. version 7.1.7.2. (Available from: http://www.diva-gis.org/) Nieto, C. 2010. Cladistic analysis of the family Baetidae (Insecta: Hovenkamp, P. 1997. Vicariance events, not areas, should be used Ephemeroptera) in South America. Systematic Entomology in biogeographical analysis. Cladistics 13:67–79. 35:512–525. 000 | The Baetodes complex C. Nieto

Nieto, C., and T. Derka. 2012. A new species of the genus cance for the ground plan of the Baetoidea. Pages 536–549 Spiritiops (Ephemeroptera, Baetidae) from the Pantepui bio- in P. Landolt and M. Sartori (editors). Ephemeroptera and geographical province. Zootaxa 3256:47–63. Plecoptera. Biology–Ecology–Systematics. Mauron + Tinguely Ortíz-Jaureguizar, E., and G. A. Cladera. 2006. Paleoenvironmen- and Lachat, SA, Fribourg, Switzerland. tal evolution of southern South America during the Cenozoic. Stöckli, R. 2015. NASA’s Earth Observatory Group. Images, us- Journal of Arid Environments 66:498–532. ing data provided by the MODIS Land Science Team. Na- Roback, S. S. 1966. The Catherwood Foundation Peruvian- tional Aeronautics and Space Administration, Washington, DC. Amazon Expedition. VI: Ephemeroptera nymphs. Monographs of (Available from: http://neo.sci.gsfc.nasa.gov) the Academy of Natural Sciences of Philadelphia 14:129–199. Waltz, R. D., and W. P. McCafferty. 1985. Moribaetis: a new Salles, F. F., J. D. Batista, and H. R. S. Cabette. 2004. Baetidae genus of Neotropical Baetidae (Ephemeroptera). Proceed- (Insecta: Ephemeroptera) de nova xavantina, Mato Grosso, ings of the Entomological Society of Washington 87:239– Brasil: novos registros e descrição de uma nova espécie de 251. Cloeodes Traver. Biota Neotropica 4(2):1–8. Wang,T.Q.,andW.P.McCafferty. 1996. New diagnostic char- Staniczek, A. H. 1997. The morphology of Siphlanenigma janae acters for the mayfly family Baetidae (Ephemeroptera). Ento- Penniket (Ephemeroptera, Siphlaenigmatidae), and its signifi- mological News 107:207–212.

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