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Schriftleitung
HANS-PETER TSCHORSNIG
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Erscheinungsdatum 30. April 2016
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Stuttgarter Beitrage zur Naturkunde A, Neue Serie 9, 2016
Inhalt / Contents
SALVADOR, R. B. & SIMONE, L. R. L.
SIMONE, L. R. L. &
SALVADOR, R. B.
SCHMALFUSS, H. MALZACHER, P. & STANICZEK, A. H.
MALZACHER, P.
BAEHR, H.
ASSING, V.
HERRMANN, A. & HAVA, J. GRIMM, R.
GRIMM, R.
SCHAWALLER, W.
SCHAWALLER, W.
SCHAWALLER, W.
KRATOCHWIL, A.
BARTSCH, D.
SCHAFER, F. & SCHMITZ, G.
A new species of Kora from Bahia, Brazil (Gastropoda: Pulmonata: Orthalicoidea), with an emended diagnosis of the genus
Taxonomical study on a sample of land snails from Nanuque (Minas Gerais, Brazil), with descriptions of three new species
Two new species and new records of terrestrial isopods (Isopoda: Oniscidea) from Oman
Two new genera of Caeninae (Ephemeroptera: Caenidae), with a cladistic analysis of the tribe Clypeocaenini
Two new Caenis species from north-eastern China (Insecta: Ephemeroptera)
New tachyine species from the Oriental Region (Coleoptera: Carabidae: Bembidiini: Tachyina)
A revision of Zyras Stephens sensu strictu of China, Taiwan, and Hong Kong, with records and (re-)descriptions of some species from other regions (Coleoptera: Staphylinidae: Aleocharinae: Lomechusini)
Attagenus (s. str.) pseudorobustior n. sp. (Coleoptera: Dermestidae: Attagenini) from Namibia
A new species of Promethis Pascoe from West Papua with unusual head armature (Coleoptera: Tenebrionidae: Cnodalonini)
New and little known species of Tenebrionidae (Coleoptera) from Borneo (6)
The genus Menimus Sharp (Coleoptera: Tenebrionidae: Gnathidiini) in India, with descriptions of two new species
Leiochrinini (Coleoptera: Tenebrionidae: Diaperinae) from north-eastern India and China, with descriptions of six new species
New species of the genus Menimus Sharp (Coleoptera: Tenebrionidae: Gnathidiini) from Peninsular Malaysia and adjacent southern Thailand
Review of the Icelandic bee fauna (Hymenoptera: Apoidea: Anthophila)
Revisionary checklist of the southern African Osminiini (Lepidoptera: Sesiidae)
Skull identification key for Central European shorebirds (Aves: Charadriiformes: Scolopaci and Charadrii)
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41
63 71
87
177
181
185 191
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Stuttgarter Beiträge zur Naturkunde A, Neue Serie 9: 1-7; Stuttgart, 30.1V.2016. DOI: 10.18476/sbna.v9.al
A new species of Kora from Bahia, Brazil (Gastropoda: Pulmonata:
Orthalicoidea), with an emended diagnosis of the genus
RopriGo B. SALVADOR & Luiz RICARDO L. SIMONE
Abstract
A new species of land snail was recently found in the municipalities of Carinhanha, Serra do Ramalho and Coribe, western Bahia state, Brazil. It is described herein as Kora rupestris n. sp. and can be easily recognized from its congeners by its usually narrower shell and aperture, and especially by its protoconch sculpture pattern. In light of this new discovery, new emended diagnosis and description are provided for the genus Kora, and two taxa pre- viously described under this genus are excluded, being transferred to the genus Drymaeus [D. iracema (Simone, 2015) n. comb. and D. terreus (Simone, 2015) n. comb.]. The region where the new species was found consists of a contact zone of the Caatinga and Cerrado biomes. Such regions are proving to be quite diverse and a more thorough knowledge of their fauna is of utmost importance for future conservation efforts.
Key words: Bulimulidae, Carinhanha, Caatinga, Cerrado, Kora rupestris new species.
Zusammenfassung
Eine neue Landschneckenart aus Brasilien wurde kürzlich in den Gebieten von Carinhanha, Serra do Ramalho and Coribe im Westen des Bundesstaates Bahia festgestellt. Sie wird hier als Kora rupestris n. sp. beschrieben und ist von anderen Arten der Gattung Kora leicht zu unterscheiden durch ihre üblicherweise schmalere Schale und Mündung, vor allem aber durch die Skulptierung des Protoconchs. Aufgrund dieser neu entdeckten Art wird eine aktualisierte Diagnose und Beschreibung der Gattung Kora gegeben und zwei zuvor in dieser Gattung beschrie- bene Arten werden in die Gattung Drymaeus gestellt [D. iracema (Simone, 2015) n. comb. und D. terreus (Simone, 2015) n. comb.]. Die Region, in der die neue Art gefunden worden ist, stellt eine Berührungszone der Caatinga- und Cerrado-Biome dar. Solche Kontaktzonen sind sehr artenreich und eine bessere Kenntnis ihrer Fauna ist von hoher Bedeutung für künftige Naturschutzbestrebungen.
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1 Introduction
Kora Simone, 2012 (Gastropoda: Pulmonata: Orthal- icoidea) is an endemic central-eastern Brazilian genus of pulmonate land snails, occurring in the Caatinga biome of the states of Bahia and Minas Gerais (SIMoNnE 2015). The genus currently includes four species: Kora coral- lina Simone, 2012 (the type species), K. iracema Simone 2015, Kora nigra Simone, 2015 and K. terrea Simone, 2015. New material from Bahia (and re-examination of the material studied by Simone 2012, 2015) has brought to light yet another species, which is described below. More- over, in light of this new discovery, an emended diagnosis for the genus Kora is provided herein.
Acknowledgements
We are deeply grateful to Jost CoLtro (Femorale) and MARIA E. BicHuETTE (Universidade Federal de Sao Carlos, Brazil) and their respective teams for the donation of the material studied
here; to DANIEL C. CAVALLARI (MZSP) for locating the material of the J. Vaz collection; to KARIN WoLF-SCHWENNINGER (SMNS) for her help with ESEM images; to Francisco J. BORRERO and an anonymous reviewer for their helpful comments on the manuscript. This work was partly supported by a doctorate grant from CNPq (Conselho Nacional de Desenvolvimento Cientifico e Tecnolögico, Brazil; proc. #245575/2012-0) to R. B.S.
2 Material and methods
The material of the new species described herein was recov- ered in the municipalities of Carinhanha and Serra do Ramalho, western Bahia state, Brazil (Fig. 1), and is listed and discussed below. For detailed examination of the protoconch, uncoated samples were mounted on stubs and observed under an Environ- mental Scanning Electron Microscope (ESEM) in the SMNS.
Furthermore, in order to present emended generic diag- nosis and description of the genus Kora, all available material described in the genus Kora was analyzed (see SIMONE 2012, 2015 for more detailed information on these lots): Kora
2 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
BAHIA
Serra do
Ramalho ea
© eo Carinhanha
Fig. 1. Map showing the presently known distribution of Kora rupestris n.sp., the municipalities of Carinhanha (type local- ity), Serra do Ramalho and Coribe. — Abbreviations of neigh- boring states: MA, Maranhäo; PI, Piaui; PE, Pernambuco; AL, Alagoas; SE, Sergipe; TO, Tocantins; GO, Goias; MG, Minas Gerais; ES, Espirito Santo.
corallina. MZSP 103910 (holotype; Bahia state, Santa Maria da Vitoria); MZSP 103911 (paratype; from type locality); MZSP 103912 (paratype; from type locality), MZSP 103913 (paratypes; 32 shells; from type locality); MZSP 104033 (>50 shells; from type locality); MNHN IM-2012-37362 (paratypes; 2 shells; from type locality); MNRJ 30377 (paratypes; 2 shells; from type locality); USNM 1157009 (paratypes; 2 shells; from type locality). — Kora iracema: MZSP 104964 (holotype; Bahia state, Sao Desidério). — Kora nigra: MZSP 106232 (holotype; Bahia state, Serra do Ramalho); MZSP 104839 (paratypes; 5 shells; unknown provenance, but likely from type locality); MZSP 106241 (paratype; from type locality); MZSP 106250 (para- types; 2 shells; from type locality). - Kora terrea: MZSP 106215 (holotype; Minas Gerais state, Presidente Olegario).
Abbreviations
D shell greatest width d aperture width H shell length h aperture height S spire (excluding aperture) S: spire (excluding body whorl) Acronyms of Institutions MNHN Museé National d’Histoire Naturelle (Paris, France) MNRJ Museu Nacional da Universidade Federal do Rio
de Janeiro (Rio de Janeiro, Brazil)
Neue Serie 9
MZSP Museu de Zoologia da Universidade de Sao Paulo (Sao Paulo, Brazil)
SMNS Staatliches Museum für Naturkunde Stuttgart (Stuttgart, Germany)
USNM National Museum of Natural History, Smith-
sonian Institution (Washington D.C., USA).
3 Systematics
Superfamily Orthalicoidea Family Bulimulidae Genus Kora Simone, 2012
Type species: Kora corallina Simone, 2012; Recent, Brazil.
Included taxa: Kora corallina Simone, 2012 (Figs. 2, 12-14); K. nigra Simone, 2015 (Fig. 3), K. rupestris n.sp. (Figs. 4-11, 15-17).
Original diagnosis (SIMONE 2012: 432)
“Outline fusiform: spire tall, somewhat turriform. Pro- toconch simple, paucispiral, ornamented by scanty axial cords. Umbilicus narrow. Peristome somewhat away from longitudinal axis of spire. Peristome deflected. Inner lip with strong, oblique tooth in middle level.”
Emended diagnosis
Shell conical-fusiform. Color initially lighter, becom- ing darker towards aperture; light-colored subsutural spi- ral band always present. Protoconch (~2 whorls) initially smooth, becoming sculptured by well-marked sinuous axial ribs later on, which gradually increase in strength until they become similar to teleoconch sculpture; propor- tion of whorls with and without sculpture pattern varies interspecifically and ribs of “transitional” strength might not be present. Transition from proto- to teleoconch not clearly marked. Teleoconch sculptured by sinuous proso- cline axial ribs. Aperture broad, slightly expanded laterally, away from the shell axis. Peristome reflexed. Columellar lamella oblique. Umbilicus narrow, but well-marked.
Discussion
The set of diagnostic features of the genus Kora allows a clear distinction from other orthalicoid genera. The main characters are analyzed below; the most important of which is the protoconch sculpture pattern.
Shell: The shell seems to be conservative within the genus, with all species displaying a conical-fusiform Shell, differing only in its proportions and absolute size. As already pointed out by Simone (2012), the degree of intraspecific conchological variation of Kora species is not high (K. corallina is known from several dozens of specimens): the variation in size is small, but shape can go from narrower to broader shells, the latter sometimes also with a broader aperture.
SALVADOR & SIMONE, NEW SPECIES OF KORA FROM BAHIA, BRAZIL 3
Color: Both K. corallina and K. rupestris n. sp. have initially light-colored shells, which become clearly darker towards the aperture. Kora nigra is more uniformly colored, although the same pattern may be observed. Var- iation in overall coloration and color pattern is seen only interspecifically in Kora (at least in the presently availa- ble material). A great amount of variation (both inter- and intraspecific) is common in other orthalicoid genera, such as, for instance, Drymaeus Albers, 1850 and Leiostra- cus Albers, 1850, but is minimal in others, such as Ple- kocheilus Guilding, 1828 and Dryptus Albers, 1860 (e. g., SIMONE 2006; BREURE & MOGOLLON AvıLA 2010; BORRERO & BREURE 2011; SALVADOR & CAVALLARI 2013; SALVADOR et al. 2015). A constant feature of the genus seems to be a lighter-colored spiral band right below the suture.
Aperture: A reflexed peristome is found in all presently-known species of Kora. This is not exclusive of Kora and can be seen in many orthalicoid genera (e. g., SIMONE 2006), although this feature is not necessarily con- stant throughout each genus.
Apertural barriers: The columellar lamella seen in Kora is a recurrent feature ın orthalicoid families (and its presence/absence may vary within some genera) and might not be very useful for diagnosis and phyloge- netic analyses. As pointed out by Simone (2012), other Brazilian orthalicoid genera with a similar lamella are Dryptus, Plekocheilus and Eudolichotis Pilsbry, 1896. A similar lamella is known in the family at least since the Late Paleocene or Early Eocene, as shown by the fossil genus /taborahia Maury, 1935, from Rio de Janeiro state (SALVADOR & SIMONE 2013).
Protoconch: The most characteristic feature of the genus Kora is the protoconch. The protoconch (~ 2 whorls) is initially smooth, becoming sculptured by well-marked sinuous axial ribs later on, which gradually increase in strength until they become similar to teleoconch sculp- ture. The exact numbers of smooth and sculptured whorls varies interspecifically (Figs. 12-17). The transition from proto- to teleoconch is weakly marked.
On the one hand, both K. corallina (Figs. 12-14) and K. nigra have the first ~ 4 whorl smooth; the next ~ 1% whorl is sculptured by faint axial striae; close to the final ~ '/4 whorl, these striae become more strongly marked, equal to the teleoconch’s ribs. On the other hand, K. rupestris n. sp. (Figs. 15-17) has the first 1/2 to 1% whorl smooth; the last % to '4 whorl has the same strong axial ribs as the teleoconch. This means that the “transitional” faintly stri- ated region is not usually present in K. rupestris n. sp., but a single specimen shows very faint axial striae immedi- ately before the onset of the thicker teleoconch-like ribs.
This axial sculpture pattern of the protoconch is sim- ilar to that of the genus Naesiotus Albers, 1850 and Rab- dotus Albers, 1850, with its sinuous and well-marked axial ribs (BREURE 1978, 1979; BREURE & Coppois 1978). Never-
theless, these two genera show the entire protoconch cov- ered in ribs, contrary to Kora, which has a smooth region before the beginning of the ribbed area. Finally, as pointed out by Simone (2012), Neopetraeus von Martens, 1885 also has axial riblets similar to Kora, but its protoconch is also sculptured by fine spiral striae, giving it a reticulate appearance (BREURE 1979).
In addition, the protoconchs of Naesiotus spp. exhibit a lower number of axial ribs than those of Kora; the number of axial ribs in the protoconch of Rabdotus spp. is intermedi- ate between those of species of Naesiotus and Kora (BREURE 1978, 1979). The pattern of the protoconch sculpture, closely reminiscent of Naesiotus and Rabdotus, might indicate the placement of the genus Kora in the family Bulimulidae (as per BREURE & ROMERO 2012) instead of the originally pro- posed allocation in Orthalicidae (Simone 2012). Protoconch sculpture tends to be a good indicative of the relationship between taxa, but the family Bulimulidae also counts with genera with very distinct protoconch patterns, such as Dry- maeus (BREURE 1978, 1979; BREURE & ROMERO 2012). As such, the importance of this character in the classification of the Orthalicoidea remains somewhat uncertain.
Finally, the protoconchs of Kora iracema Simone, 2015 and K. terrea Simone, 2015 exhibit a completely reticulated sculpture pattern, thus lacking the characteris- tic protoconch of Kora. This conformation is characteris- tic of the genus Drymaeus (BREURE 1979). Moreover, their overall shell shape, aperture shape, lack of a strong colu- mellar fold, and greatly reflexed peristome are also befit- ting of the genus (BREURE 1978, 1979; SımonE 2006). As such, both species are here transferred to this genus as Drymaeus iracema (Simone, 2015) n.comb. and D. ter- reus (Simone, 2015) n. comb.
Kora rupestris n. sp. (Figs. 4-11, 15-17)
Kora corallina: Simone 2015 [in part]: 53 (figs. 14, 22).
Holotype: MZSP 121416 (A. Biancur col., VIII.2012; Figs. 4-7, 15-17).
Paratypes: MZSP 121441 (2 shells, from type locality; A. BIANcHI col., VIII.2012; Figs. 8-9).
Additional material: MZSP 106230 (1 shell; Serra do Ramalho municipality, Gruna das Trés Cobras; 14°19'S 43°47'W, alt. 440m; M.E. BicHUFTTE col., 10.1X.2008; Fig. 10). MZSP 107996 (1 shell; Coribe municipality; J. Vaz collection; M. U. T. Ropricues col., 1.1980; Fig. 11).
Type locality: Brazil, Bahia, Carinhanha municipality (city center coordinates: 14°18'18"S 43°45'54"W), under rocks on Canabrava Hill. Unfortunately, more precise locality data, such as coordinates and altitude, were not recorded by the collector.
Etymology
From the Latin “rupestris”, meaning “of rock”, “montane”. Reference to the rocky mount where the types were collected.
4 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
Neue Serie 9
Figs. 2-11. Kora spp. from Bahia state, apertural (2-4, 8-11), umbilical (7), lateral (5), and dorsal (6) views. — 2. K. corallina (holo- type, MZSP 103910; H=43.4mm). 3. K. nigra (holotype, MZSP 106232; H=30.1 mm). — 4-7. K. rupestris n. sp. (holotype, MZSP 121416, H=37.7 mm). — 8. K. rupestris n. sp. (paratype #1, MZSP 121441; H=35.5 mm). 9. K. rupestris n. sp. (paratype #2, MZSP 121441; H=33.3 mm). 10. K. rupestris n. sp. (MZSP 106230; H=34.2 mm). 11. K. rupestris n.sp., apertural view (MZSP 107996; H=35.6 mm).
Distribution
Known only from the type locality and the neighbor- ing municipalities of Serra do Ramalho to the north and Coribe to the west (Fig. 1).
Habitat
Caatinga shrublands.
Diagnosis
Shell slender. Shell color comprised of light tones of brown. Protoconch sculpture pattern: first 1% to 1% whorl
smooth; last % to Y% whorl sculptured by well-marked axial striae. Aperture narrow.
Description
Shell large (~ 35 mm), conical-fusiform, slender; great- est width on body whorl; shell width ~7/s shell length (Figs. 4-6, 8-11). Shell color light brown; last whorl increasingly darker; narrow stripe immediately below the suture slightly lighter than remainder of whorl (Figs. 4—6). Peristome white. Spire angle usually ca. 45°, but up to ca. 50° in broader specimens. Protoconch (~2 whorls; Figs. 15-17) rounded; first 172 to 1% whorl smooth; last
SALVADOR & SIMONE, NEW SPECIES OF KORA FROM BAHIA, BRAZIL 5
Figs. 12-17. Kora spp. from Bahia state, lateral (12-13, 15-16), and apical (14, 17) views. — 12. K. corallina, proto- and teleoconch sculpture (paratype, MZSP 103913). 13. Same, in higher magnification, showing the transition from faint to more marked sculp- ture. 14. Same, showing the transition from smooth to faintly sculptured protoconch. 15. K. rupestris n. sp., lateral view of proto- and teleoconch sculpture (holotype, MZSP 121416). 16. Same, in higher magnification, showing the transition from smooth to sculptured protoconch. 17. Same, showing the almost entirely smooth protoconch. — Scales: 1 mm (12, 15), 0.5mm (13-14, 16-17).
Y2 to Ya whorl sculptured by well-marked axial striae (same as teleoconch); transition to teleoconch not clearly marked, prosocline, simple. Teleoconch sculptured by sin- uous prosocline axial striae (Fig. 15). Whorl profile lightly convex. Suture well-marked, but not deep. Body whorl almost % shell length. Aperture oval, prosocline, propor- tionately large, slightly expanded laterally away from shell axis; aperture ~ % shell length, -% shell width. Columel- lar lamella well-marked, variable in size, oblique to shell axis (Figs. 4, 8-11). Peristome greatly reflexed. Umbilicus narrow (Fig. 7).
Measurements (in mm). Holotype: 6 whorls; H = 37.7; D=16.3; S=19.4, S’=13.7; h=18.2; d=11.4. — Para- type #1: 6% whorls; H=35.5; D=15.3; S=18.6; S’ =12.4; h=16.5; d=10.7. — Paratype #2: 6% whorls; H=33.3; D=14.8, S=171; S’=12.2; h=16.5; d=10.2. — MZSP 106230: 6% whorls; H=34.2; D=15.9; S=17.9; S’ =12.0; h=15.9; d=11.1. — MZSP 107996: 6% whorls; H=35.6; D=18.2; S=18.3; S =13.1; h=17.6; d=12.1.
Differential diagnosis
Kora rupestris n. sp. can be easily diagnosed by its dif- ferent protoconch sculpture pattern (see the discussion of
the genus, above, for the comparison with K. corallina and K. nigra). Moreover, K. rupestris n. sp. (Figs. 4, 8-11) has a more slender shell and spire than K. corallina (Fig. 2) and X. nigra (Fig. 3), which also results in a slightly nar- rower aperture; both X. corallina are and K. nigra are very broad, as also reflected in their greater spire angle values of 45-50° (K. corallina) and 60-65° (K. nigra). However, there are more slender specimens of K. coral- lina, which present the same shell length/width ratio as K. rupestris n. sp. Such specimens may be diagnosed by size (besides the protoconch), as the shell of K. rupestris n. sp. is smaller than that of K. corallina, having approxi- mately % of the shell length of the latter, with no overlap in their size ranges. Kora rupestris n. sp. is slightly larger than K. nigra, although some doubtful specimens of the latter (MZSP 104839) are in the upper size range of K. corallina. Furthermore, K. rupestris n.sp. shows % to % less whorl in the teleoconch, when compared to K. cor- allina, although it has the same whorl count as K. nigra. Finally, the color pattern of K. rupestris n. sp. (Fig. 4) is similar in nature to that of K. corallina (Fig. 2), but it is usually comprised of overall lighter tones of brown; K. nigra has a more homogenous dark coloration (Fig. 3).
6 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
The single specimen (MZSP 106230) recovered in a cavern and identified by Simone (2015: figs. 14, 22) as K. corallina, in fact belongs to K. rupestris n.sp. It agrees with K. rupestris n.sp. in size, overall shape and, more importantly, the protoconch sculpture pattern. This sin- gle specimen is slightly broader than the type specimens of K. rupestris n.sp., with a narrower umbilicus, propor- tionately smaller aperture and displaying a stronger white parietal callus. Nevertheless, this same kind and extent of variation is seen in the type series of K. corallina, and is thus considered here intraspecific variation of K. rupes- tris N. sp.
4 Concluding remarks
Carinhanha and Serra do Ramalho municipalities, the known distribution of Kora rupestris, are located in western Bahia state (Fig. 1), on the west banks of the Sao Francisco River. It is an interesting region, consisting of a contact zone of the Caatinga and Cerrado biomes, with the hypoxerophytic forests of the former and stretches of the open deciduous forest of the latter (SA et al. 2003). The cli- mate 1s considered tropical dry (semiarid), with a dry win- ter season and rain concentrated on the end of spring and beginning of summer. It is a limestone-rich region, which seems to be favorable for land snails, perhaps especially for those with sturdy shells (e. g., SALVADOR & SIMONE 2014).
The Caatinga biome, despite being usually considered of low biodiversity levels, is now revealing an amazing array of species, across all main terrestrial and fresh- water taxa and including several endemic species (e. g., TABARELLI & SiLvA 2002). Nevertheless, there are still proportionally few works on this fauna, which is often not included in conservation measures (e. g., SILVA et al. 2003). This is especially troubling, since circa 20% of the Caatinga is already considered degraded (SA et al. 2003) and continental mollusks are considered the most imperiled group of animals, with the highest extinction rates (LYDEARD et al. 2004; REGNIER et al. 2008). The more arid habitats are normally seen as an adverse place for land snails (which depend so much on preserving water), but it appears that the Caatinga may still hide many new snails (e.g., SALVADOR & SIMONE 2014; SımonE 2015). This is especially true due to the limestone-rich environments of this biome, which allow land snails to thrive, but which are now threatened by mining activities (SIMONE 2015). More- over, the specimens of Kora nigra and Drymaeus iracema (as well as a single specimen of Kora rupestris) were found in caves (Simone 2015), which are fragile ecosystems and thus face their own set of environmental problems (SIMONE 2013; Campos-FILHo et al. 2014). A more thorough knowl- edge of this biome’s species is of utmost importance for future conservation efforts.
Neue Serie 9
5 References
BORRERO, F. J. & BREURE, A.S.H. (2011): The Amphibulimidae (Mollusca: Gastropoda: Orthalicoidea) from Colombia and adjacent areas. — Zootaxa 3054: 1-59.
BREURE, A. S. H. (1978): Notes on and descriptions of Bulimuli- dae (Mollusca, Gastropoda). — Zoologische Verhandelingen 164: 1-255.
BREURE, A. S. H. (1979): Systematics, phylogeny and zoogeogra- phy of Bulimulinae (Mollusca). — Zoologische Verhandelin- gen 168: 1-215.
BREURE, A.S. H. & Coppois, G. (1978): Notes on the genus Nae- siotus Albers, 1850 (Mollusca, Gastropoda, Bulimulidae). — Netherlands Journal of Zoology 28: 161-192.
BREURE, A. S. H. & MoGoLLön Avia, V. (2010): Well-known and little-known: miscellaneous notes on Peruvian Orthalicidae (Gastropoda, Stylommatophora). — Zoologische Mededelin- gen 84: 15-35.
BREURE, AS. H. & Romero, P. E. (2012): Support and surprises: a new molecular phylogeny of the land snail superfam- ily Orthalicoidea (Gastropoda, Stylommatophora) using a multi-locus gene analysis. — Archiv für Molluskenkunde 141: 1-20.
CAMPOS-FILHO, I. S., ARAUJO, P. B., BICHUETTE, M. E., TRAJANO, E. & Taiti, S. (2014): Terrestrial isopods (Crustacea: Isopoda: Oniscidea) from Brazilian caves. — Zoological Journal of the Linnaean Society 172: 360-425.
LYDEARD, C., CowIE, R. H., PONDER, W. F., BoGAN, A. E., BOUCHET, P., CLARK, S. A., CumMInGs, K. S., FREST, T. J., GARGOMINY, O., HERBERT, D. G., HERSHLER, R., PEREZ, K. E., ROTH, B., SEDDON, M., Strong, E.E. & Thompson, F.G. (2004): The global decline of nonmarine mollusks. — Bioscience 54: 321-330.
REGNIER, C., FONTAINE, B. & BoucHET, P. (2008): Not know- ing, not recording, not listing: numerous unnoticed mollusk extinctions. — Conservation Biology 23: 1214-1221.
SA, 1.B., RicHé, G.R. & Fortius, G. A. (2003): As paisagens e o processo de degradacäo do semi-arido nordestino. — In: SıLva, J.M.C., TABARELLI, M., Fonseca, M. T. & Lins, L. V. (eds.): Biodiversidade da Caatinga: areas e acÖes prioritarias para a conservagäo. pp. 17-36; Brasilia (Ministério do Meio Ambiente & Universidade Federal de Pernambuco).
SALVADOR, R. B. & CAVALLARI, D.C. (2013): Taxonomic revision of Leiostracus onager and Leiostracus subtuszonatus (Gas- tropoda: Pulmonata: Orthalicidae). — Journal of Conchology 41: 511-518.
SALVADOR, R. B. & Simone, L.R. L. (2013): Taxonomic revision of the fossil pulmonate mollusks of Itaborai Basin (Pale- ocene), Brazil. — Papéis Avulsos de Zoologia 53: 5—46.
SALVADOR, R. B. & SIMONE, L. R. L. (2014): New species of Cyclo- dontina from Bahia, Brazil (Gastropoda, Pulmonata, Odon- tostomidae). — Iheringia, Série Zoologia 104: 484—487.
SALVADOR, R.B., CAVALLARI, D.C. & Simone, L.R.L. (2015): Taxonomical study on a sample of land snails from south- eastern Tocantins state, Brazil, with description of a new species. — Journal of Conchology 42: 67-78.
SıLva, J.M.C., TABARELLI, M., Fonseca, M.T. & Lins, L. V. (2003): Biodiversidade da Caatinga: areas e acöes priori- tarias para a conservacäo, 382 pp.; Brasilia (Ministério do Meio Ambiente & Universidade Federal de Pernambuco).
SIMONE, L. R. L. (2006): Land and freshwater mollusks of Brazil, 390 pp.; Sao Paulo (Editora Grafica Bernardi & Fundacäo de Amparo a Pesquisa do Estado de Säo Paulo).
SIMONE, L. R. L. (2012): Taxonomical study on a sample of pul- monates from Santa Maria da Vitoria, Bahia, Brazil, with
SALVADOR & SIMONE, NEW SPECIES OF KORA FROM BAHIA, BRAZIL i
description of a new genus and four new species (Mollusca: TABARELLI, M. & Siva, J.M.C. (2002): Areas e agöes priori-
Orthalicidae and Megalobulimidae). — Papéis Avulsos de tarias para a conservacao, utilizacäo sustentavel e reparticäo
Zoologia 52: 431—439. de beneficios da biodiversidade do bioma Caatinga. — In: SIMONE, L.R.L. (2013): Habeas, a new genus of Diplommati- ARAUJO, E. L., Moura, A.N., Sampaio, E. V.S. B., GESTINARI,
nidae from Central Bahia, Brazil (Caenogastropoda), with L.M.S. & Carneiro, J.M.T. (eds.): Biodiversidade, Con-
description of three new species. — Journal of Conchology servacäo e Uso Sustentavel da Flora do Brasil, pp. 47-52;
41: 519-525. Recife (Sociedade Botanica do Brasil & Universidade Fed- SIMONE, L. R. L. (2015): Three new species of Kora (Pulmonata, eral Rural de Pernambuco).
Orthalicidae) from Bahia and Minas Gerais, Brazil. — Jour- nal of Conchology 42: 51-56.
Authors’ addresses:
RopriGO BRINCALEPE SALVADOR (corresponding author), Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191, Stutt- gart, Germany; e-mail: salvador.rodrigo.b@gmail.com
Luiz Ricarpo Lopes SIMONE, Museu de Zoologia da Universidade de Säo Paulo, Avenida Nazare 481, 04218970 Sao Paulo, Brazil; e-mail: Irsimone@usp.br
Manuscript received: 16.VIII.2015, accepted: 14.X.2015.
BHL in An
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Stuttgarter Beiträge zur Naturkunde A, Neue Serie 9: 9-30; Stuttgart, 30.1V.2016. DOI: 10.18476/sbna.v9.a2
Taxonomical study on a sample of land snails from Nanuque (Minas Gerais, Brazil), with descriptions of three new species
Luiz RICARDO L. SIMONE & RODRIGO B. SALVADOR
Abstract
A sample of land snails was recently collected in a fragment of Atlantic rainforest, in the vicinities of the city of Nanuque (north of Minas Gerais state, Brazil), totaling 15 species. The following new species are herein described: Leiostracus carnavalescus n. sp. and Rhinus botocudus n. sp. (Bulimulidae), and Obeliscus boitata n. sp. (Subulin- idae), the first two accompanied by anatomical descriptions. Moreover, the geographical ranges of some species are extended to Minas Gerais: Auris bilabiata, Bahiensis cf. bahiensis, Cyclopomops moricandi, Dysopeas muibum, Helicina boettgeri, Helicina variabilis, Prohappia besckei, and Rectartemon piquetensis. The discovery of new spe- cies in such a small forest fragment is a clear reminder of how little the Brazilian terrestrial snail fauna is known. It also points to the fact that these few remaining forest fragments may house many new and possibly endemic spe- cies and should, therefore, be properly preserved.
Key words: Atlantic Forest, Caenogastropoda, Gastropoda, Neritimorpha, Pulmonata, Stylommatophora.
Zusammenfassung
Fünfzehn Arten Landschnecken wurden im Mai 2012 in einem atlantischen Regenwald-Fragment in der Umge- bung der Stadt Nanuque im Norden des Bundesstaates Minas Gerais in Brasilien gesammelt. Die folgenden drei neuen Arten werden beschrieben: Leiostracus carnavalescus n.sp. und Rhinus botocudus n.sp. aus der Familie Bulimulidae sowie Obeliscus boitata n.sp. aus der Familie Subulinidae. In den ersteren beiden Beschreibungen werden auch ausführliche anatomische Merkmale berücksichtigt. Die folgenden Arten werden zum ersten Mal für Minas Gerais nachgewiesen: Auris bilabiata, Bahiensis cf. bahiensis, Cyclopomops moricandi, Dysopeas muibum, Helicina boettgeri, Helicina variabilis, Prohappia besckei und Rectartemon piquetensis. Die Entdeckung der drei für die Wissenschaft neuen Arten in dem kleinen Waldfragment ist ein deutlicher Hinweis darauf, wie gering die Kenntnis der brasilianischen Landschneckenfauna ist, und wie wichtig der Schutz auch kleiner Waldreste wäre, die viele neue und möglicherweise endemische Arten enthalten könnten.
Contents IE vlntroduetion.. aM en En EEE ZU U. Be ee Aas A DE N ER OA EEE m Bo BEE, DE A ays OB FERN LEN RN 9 2.7 HM atelsals:and IIe LO Cs: Int nt. ee Se en Meter Seca TE TI ran ar ee eee eer Pitan war ks eat 9 Sham SV SLCIITALIC Se Fr LEE Are Amber ee Meinen) Arme Werte en ich an Aare Jame Warten Ar ce Bart rien Are A 12 As MIDIS CUSSION ERS Te Na ENTE NE TEE BUCS eR RUE TB ne AN 29 I ERTEILEN CORA Rcd ol Bigs CERIN Tr En ad BEE. eect UO Bed A «CTR Ener Rd 3 ee RR LER Ihre 29
1 Introduction
The southeastern region of Brazil is historically the most explored (and exploited) in the country: its natural covering was fiercely degraded through the few centuries of the country’s history. However, many new discoveries still wait in the remnants of Atlantic rainforest, especially regarding the scarcely studied terrestrial molluscan fauna. An expedition in May 2012 by shell collector Josk CoLtro Jr. and his team to the previously unsampled region of Nanuque, in northern Minas Gerais state, recovered many land snails. Part of this material was donated to the col- lection of the Museu de Zoologia da Universidade de Sao Paulo (MZSP, Sao Paulo, Brazil) and is studied herein. As a testament to the incomplete knowledge of the Brazil- ian fauna, some of the animals recovered are new species. This paper presents formal descriptions of these taxa and
extends the geographical distributions of some other pre- viously known species.
Acknowledgements
We are deeply grateful to Jost Cottro Jr. (Femorale) for pro- viding the material studied here; to LARA GuIMARAES (MZSP) for helping with SEM examination; to BARBARA M. TOMOTANI (Netherlands Institute of Ecology, Wageningen, The Nether- lands), CARLO M. Cunna (Academy of Natural Sciences of Phil- adelphia, USA) and DAnIEL ABBATE (MZSP) for some of the photos used here (BMT: photos of Leiostracus from Mantena; CMC: photos of the living animals; DA: photos of Helicina boettgeri and Bahiensis).
2 Materials and methods
All the specimens collected are land snails, mainly stylom- matophoran pulmonates, but two neritimorph species and one
10 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
caenogastropod are also present. Most species are only repre- sented by dry shells, usually in a good state of preservation; only four were captured alive: Helicina variabilis, Leiostracus per- lucidus, Leiostracus carnavalescus n.sp. and Rhinus botocu- dus n.sp. All the animals were collected in the vicinities of the city of Nanuque (17°50'51"S 40°22'47"W, ~ 120m of elevation; Fig. 1), in the north of Minas Gerais state, very close to the bor- ders with the states of Bahia and Espirito Santo. The locality is a very small fragment of Atlantic rainforest (area estimated at ca. 0.15 km?), close to the Mucuri River, surrounded by tomato crops and thus highly subject to anthropic action and degradation.
The animals collected alive were preserved in 70% etha- nol. Specimens were extracted from their shells and dissected (immersed in ethanol) by standard techniques under a stereo- microscope. Digital photographs were obtained for most dissec- tion steps, as well as drawings with the aid of a camera lucida. Measurements were made with a digital caliper or with the Zeiss ZEN 2011 software. The radulae were mounted on stubs, coated with a gold-palladium alloy and examined by scanning elec- tronic microscopy (SEM) in the Laboratorio de Microscopia Eletronica of MZSP.
Identification was conducted based on the catalogue by SIMONE (2006), the original descriptions and additional material housed in the collection of the MZSP. Measurements were made
BRAZIL
MINAS GERAIS
Neue Serie 9
with a digital caliper or with the aid of the Zeiss Axiovision SE64 Rel 4.8 imaging software. All specimens are deposited in the malacological collection of the MZSP. The list of examined material can be found together with the descriptions of the new species and, for the other species, on Tab. 1. Interestingly, spec- imens of the new Leiostracus species were found in the MZSP collection (from the municipalities of Mantena, Minas Gerais state, and Pinheiros and Sooretama, Espirito Santo state; Fig. 1) and were also included in the analysis.
Abbreviations Shell measurements
D shell greatest width (diameter)
d aperture width
H shell height
h aperture height
S spire height (excluding aperture)
Ss’ spire height (excluding body whorl) Anatomical characters
aa anterior aorta
ac albumen chamber
ad albumen gland duct
Fig. 1. Map showing the municipality of Nanuque, where the specimens studied here were collected, and the other municipalities from where the specimens of Leiostracus carnavalescus n.sp. come from: (1) Mantena; (2) Pinhei- ros; (3) Sooretama. — Abbreviations of the neighboring states: BA, Bahia; GO, Goias; ES, Espirito Santo; RJ, Rio de Janeiro; SP, Sao Paulo.
SIMONE & SALVADOR, TAXONOMICAL STUDY ON A SAMPLE OF LAND SNAILS FROM NANUQUE 11 Tab. 1. Other species occurring in the present material from Nanuque, accompanied by data on the material analyzed, mean mea- surements (of adult specimens) and previously known geographic distribution (sensu Morretes 1949, SIMONE 2006). *Record from RoBINSON & SLAPCINSKY (2005). — Abbreviations: Material: sh., shell; spc., specimen. Brazilian states: BA, Bahia; CE, Ceara; ES,
Espirito Santo; MG, Minas Gerais; PB, Paraiba; PE, Pernambuco; RJ, Rio de Janeiro; RN, Rio Grande do Norte; SC, Santa Cata- rina; SE, Sergipe; SP, Sao Paulo.
(Broderip & Sowerby, 1829)
(d’Orbigny, 1835) Brazil, Bolivia, Uruguay (Spix, 1827) 106161 (1 spc.)
Wagner, 1910 106167 (6 sh.)
Wagner, 1827 (4 spc.), 106173 (1 sh.)
(Pfeiffer, 1852)
Bahiensis cf. bahiensis Odontostomidae 106165 (1 sh.) H=16.2, D=5.1 Brazil (BA, RJ) (Moricand, 1833)
Rectartemon piquetensis Streptaxidae 106156 (9 sh.) H=20.3, D=31.6 Brazil (BA, SP) (Pilsbry, 1930)
Megalobulimus bronni Strophocheilidae 106154 (1 sh.) H=91.3, D=56.0 Brazil (MG, ES, SP) (Pfeiffer, 1847)
Dysopeas muibum Subulinidae 106163 (1 sh.) H=5.7, D=2.4 Brazil (SP)
Marcus & Marcus, 1968
Succinea meridionalis Succineidae 106162 (1 sh.) H=8.2, D=49 Brazil (RJ) to Argentina d’Orbigny, 1837 (Patagonia)
Prohappia besckei Systrophiidae 106164 (6 sh.) H=44,D=7] Brazil (RJ, SC), Paraguay (Dunker in Pfeiffer, 1847)
ag albumen gland gd genital diverticulum
an anus gf gastric inner fold
ap genital aperture go gonad
au auricle hd hermaphrodite duct
bc bursa copulatrix in intestine
bd bursa copulatrix duct ir insertion of m4 in subradular cartilage bg buccal ganglia jw jaw
bm buccal mass ki kidney chamber
br subradular membrane kl kidney lobe
ee cerebral ganglion ml—m13 __ extrinsic and intrinsic odontophore muscles cm columellar muscle mb mantle border
cn cerebro-pedal and cerebro-pleural connectives mf mantle fold
co collar vessel mj jaw and peribuccal muscles
CV pulmonary (efferent) vein mo mouth
cy statocyst mu ommatophore retractor muscle
da digestive gland anterior lobe mx platform of mj in median line
dd duct to digestive gland ne nephrostome
df dorsal folds of buccal mass nr nerve ring
dg digestive gland posterior lobe oc odontophore cartilage
di diaphragm or pallial floor od odontophore
eh epiphallus of odontophore cartilage median fusion with its pair eo spermoviduct om ommatophore
es esophagus pa posterior aorta
ey eye pc pericardium
fd spermoviduct inner folds pe penis
ft foot pf penis inner folds
ga pedal gland aperture pg pedal gland
12 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
pi penis inner chamber
pm penis muscle
pn pneumostome
pp pedal ganglion
ps penis sheath
pt prostate
pu pulmonary cavity
pv pneumostome right flap
ra radula
rn radular nucleus
rs radular sac
rt rectum
sa salivary gland aperture
SC subradular cartilage
sd salivary gland duct
sg salivary gland
sp sperm groove
sr seminal receptacle
st stomach
su subesophageal ganglion
tg integument
tm tentacle retractor muscle
to tissue on radular ribbon preceding radular exposed region in buccal cavity
ua ureter aperture
ug urinary gutter
un union of mantle border with nuchal surface
up primary ureter
ur secondary ureter
ut uterus
vd vas deferens
ve ventricle
vf spermoviduct inner fold preceding vas deferens aperture
vg vagina
3 Systematics
Gastropoda Pulmonata Stylommatophora Family Bulimulidae Genus Leiostracus Albers, 1850
Leiostracus carnavalescus n. sp. (Figs. 2-27, 53-67)
Type material
Holotype: MZSP 106177 (Figs. 53-58). Paratypes: MZSP 106178 (14 specimens), 106179 (24 Shells), from type locality. J. Cotrro col., May/2012.
Type locality: Brazil. Minas Gerais: Nanuque, ~ 120 m of elevation (~ 17°51'S 40°23'W).
Additional material
Brazil. Minas Gerais: Mantena (city center coordinates: 18°46'55"S 40°58'48"W): MZSP 108010 (3 shells; ALEx B. Bopart col., Feb/1993). — Espirito Santo: Pinheiros: Veado stream (“Cörrego do Veado”): MZSP 106618 (4 specimens; FRANKLIN N. Santos col., 29/May/2011). — Sooretama: Con- torno road (“Estrada do Contorno”): MZSP 106920 (1 specimen;
Neue Serie 9
FRANKLIN N. Santos col., 14/Apr/2012). — Sooretama: Jequitiba trail (“Trilha do Jequitiba”’): MZSP 106666 (1 specimen; FRANKLIN N. Santos col., 18/Jun/2012). — Sooretama: Quirin- inho trail (“Trilha do Quirininho”): MZSP 106650 (1 specimen; FRANKLIN N. Santos col., 14/Apr/2012).
Etymology
The name is a reference to the amazing variation in shell color in this species, establishing a parallel with the Carnival festival, when people dress in colorful costumes.
Diagnosis
Distinctive color pattern, with great intraspecific var- iation. Most usual color pattern is: spire top red, reddish orange or orange; spire with one broad spiral red or red- dish brown band on middle portion of whorl, flanked by one or two lines of brownish dots; body whorl with dark brown or black spiral band on its base and red or light brown to dark brown spiral band close to the umbilicus.
Description
Shell (Figs. 53-65, 67): medium-sized (~ 25 mm), con- ical-oval; apex acuminated; greatest width on last whorl: width -' shell length. Base color white; color pattern with three main patterns (although some intraspecific variability can be found within these, mainly in color and thickness of the bands): (1) spire top red, reddish orange or orange; spire with one broad spiral red or reddish brown band on middle portion of whorl, flanked by one or two lines of brownish dots; body whorl with dark brown or black spiral band on its base and red or light brown to dark brown spiral band close to the umbilicus (Figs. 53-56, 59); (2) spire top orange; spire almost lacking white regions and with very thick orange spiral band on middle portion of whorl, flanked by lines of brownish dots; body whorl with many spiral bands below the orange one (from top to bottom: yellow band with the previously mentioned line of brownish dots superposed on it; dark brown or black band; another thick orange band; and, finally, a red band close to umbilicus) (Fig. 60); (3) spire top dark red to red- dish brown; spire almost lacking white regions and with very thick spiral dark brown to black band and, below it, a single line of brownish dots; body whorl with greater white portion on its bottom, one dark brown or black spi- ral band on its base and another of same color close to umbilicus (Fig. 61). Spire angle ~ 40°. Protoconch of 1.5 whorl, sculptured by fine axial sinuous parallel riblets in upper portion and numerous fine spiral striae in lat- eral portion (Fig. 58); transition to teleoconch clear. Tele- oconch smooth, except for growth lines. Whorl profile slightly convex. Suture well-marked. Aperture medium- sized, prosocline, oval; ~7/s shell length, ~?/s shell width. Peristome reflexed, especially on columellar region, par- tially covering umbilicus. Body whorl -' shell length. Umbilicus narrow.
SIMONE & SALVADOR, TAXONOMICAL STUDY ON A SAMPLE OF LAND SNAILS FROM NANUQUE 13
j . ey 7 a : s ‘ F R we f | # , PR. j a = | 7 A | re \ N ti 1 Frit ag k IP | 5 | W A Zu Ss + \ \ wy > | \ : r Fe x 2 :. Er rot ra a, AL | x ) a en ; is or | ‚v4 N m N f i er u 7 ‘ x a, ig if « ~~ y 7‘ N 4 4 4 Ne gl u ; 3 7 x EN 3 ) ( ( , p r ¥ RT: Ki ; f \- A A j \ E74 Ura 4 1 y a ee x 3 ie x i \ 1 : 7 F 2 Y ig N 5 3 nn N . om { T ; 5 7 ; f \
ELERERLLET
a. a | N ® A 8 0 9 Q N
BEER FE,
=P DD:
al on in! Fr ON A 4 Vu ii
Er Hu
Figs. 2-5. Leiostracus carnavalescus n.sp., SEM images of radulae. The arrows indicate the rachidian column. — 2. Whole view. 3. Detail of central region. 4. Greater detail of central region. 5. Detail of lateral region. — Scale bars: 20 um (4), 50 um (5), 100 um
(3), 300 um (2).
Head-foot (Figs. 14, 67): of normal shape. Color white, with a broad horizontal black band running continuously from frontal portion of head to middle portion of foot. Pair of ommatophores well-developed, with strong retrac- tor muscles (tm). Eyes dark. Pair of tentacles brownish, with - 2 ommatophores’ length. Columellar muscle thick, 2 whorls in length, divided into 5—6 separated bundles (Fig. 14: cm); originating in lateral regions of haemocoel and running along its ventral surface; bundles connecting with each other in insertion region on the columella.
Mantle organs (Figs. 6-9): mantle border thick, lack- ing pigments. Pneumostome protected by ventral right flap (Figs. 6, 7: pv) with - 4 of aperture length. Another similar-sized secondary fold is located on the left side of the pneumostome (Figs. 6, 7: mf). Pneumostome ~ 10 % of aperture length, bearing exclusively the air entrance and the urinary gutter (Fig. 7: ua, ug); anus is a separated aper- ture located on the right of the pneumostome (Fig. 7: an). Lung of 1.5 whorls in length, narrow and elongated. Pul- monary vessels conspicuous only in anterior quarter (Fig. 9); remaining regions almost smooth, with imbri- cated vessels of difficult visualization (Fig. 9: pu). Pulmo- nary vein (Figs. 8, 9: cv) running longitudinally between middle and right thirds of pallial cavity roof. Kidney beige, located posteriorly, occupying ~ 20 % of cavity length and ~50% of its width (details below). Rectum (rt) and ureter (ur) narrow, running along right edge.
Visceral mass (Fig. 14): about 3.5 whorls in length. Both digestive gland lobes pale brown in color. Ante-
rior lobe (Fig. 14: da) flattened, occupying ~ !/ of visceral volume, located just posteriorly to pallial cavity, contin- uous to kidney. Posterior lobe (dg) with 3 spiral whorls, occupying ~ 70 % of visceral volume. Stomach occupying ~'s of visceral volume, located between both digestive gland lobes, about half whorl posterior to pallial cavity (Fig. 14: st). Digestive tubes (described below) surround- ing anterior lobe of digestive gland. Gonad clearly multi- lobed, cream colored, occupying entire second whorl, encased in middle-right region of posterior lobe of diges- tive gland, occupying ~ '/is of visceral volume.
Circulatory and excretory systems (Figs. 8, 9): pericar- dium about twice as long as wide, located longitudinally between middle and left thirds of posterior end of pallial roof; occupying -5% of lung area. Auricle (au) located anteriorly, as a continuation of the pulmonary vein; slightly smaller than the ventricle (ve). Kidney (ki) sim- ple, entirely solid, dorsoventrally flattened; size reported above. Nephrostome small, located on anterior-left corner as the tip of a small projection of renal tissue, turned to the right (Fig. 8: ne). Primary and secondary ureter complete, closed (tubular); primary ureter lying on right edge of kid- ney, towards the posterior and right region, after forming a strong curve; afterwards running anteriorly, as secondary ureter (ur), along entire left edge of rectum. Urinary aper- ture (Fig. 7: ua) simple, turned anteriorly, located on right edge of pneumostome. Urinary gutter (Fig. 7: ug) running in front of urinary aperture, almost perpendicularly to mantle border, gradually disappearing.
14 STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
iE TARE \
Ne NN II NK N TR N R oN oo A oS x
See En aly
DENS REN SUSE
Figs. 6-13. Leiostracus carnavalescus n.sp., anatomy. — 6. Detail of dorsal half of the mantle border, in frontal view. 7. Pneumos- tome region, in ventral view (ventral wall deflected upwards). 8. Reno-pericardial region, in ventral view (primary ureter opened lon- gitudinally; the heart can be seen by transparency). 9. Pallial cavity, in inner ventral view (ventral edge of pneumostome deflected). 10. Foregut, in lateral (right) view (some of the more posterior structures were artificially separated from each other). 11. Buccal
mass, in lateral (right) view. 12. Buccal mass, in ventral view. 13. Spermoviduct, transverse section in its middle region. — Scale bars: 1mm (13), 2mm (6-12).
SIMONE & SALVADOR, TAXONOMICAL STUDY ON A SAMPLE OF LAND SNAILS FROM NANUQUE 15
Figs. 14-19. Leiostracus carnavalescus n.sp., anatomy. —14. Dissected specimen, largely in lateral (right) view (haemocoel opened longitudinally and deflected; visceral mass partially uncoiled; pallial cavity structures only partially shown; genital organs com- pletely removed). 15. Stomach as seen in situ, in ventral view (ventral wall opened longitudinally). 16. Buccal mass, in ventral view (odontophore partially removed and deflected). 17. Odontophore, in dorsal view (the most superficial layer of muscles and mem- branes was removed; the left muscles were deflected to the right). 18. Odontophore’s cartilages, in ventral view, showing the inser- tion marks of some muscles. 19. Odontophore, in dorsal view (radular sac partly opened longitudinally; m3 deflected downwards). — Scale bars: 2mm.
Neue Serie 9
STUTTGARTER BEITRAGE ZUR NATURKUNDE A
16
showing ori-
oy
Figs. 20-25. Leiostracus carnavalescus n. sp., anatomy. — 20. Odontophore, in ventral view (most muscles removed
gins of intrinsic muscles in cartilages: oc). 21. Isolated genital system, in complete dorsal view. 22. Same, detail of albumen gland
region (ag). 23. Same
with base of albumen gland (ag) removed to show connection with spermoviduct. 24. Penis opened longitudi-
2.
nally, showing inner surface in dorsal view (some of the adjacent structures are also shown). 25. Anterior region of spermoviduct (up to genital pore), in dorsal view (greatly opened longitudinally to show inner surface; some of the adjacent structures are also shown).
— Scale bars: 2mm.
SIMONE & SALVADOR, TAXONOMICAL STUDY ON A SAMPLE OF LAND SNAILS FROM NANUQUE 7
Digestive system (Figs. 10-12, 14—20): oral tube short, with thick muscular walls (Figs. 11, 12, 16: mj). Jaw simple; exposed portion horseshoe-shaped, bearing several uni- form transverse folds (Figs. 16: jw, 66); color beige, trans- lucent. Buccal mass spherical, ~'/s of haemocoel volume (Fig. 14: bm). Dorsal surface of oral cavity with well-devel- oped pair of dorsal folds (Fig. 16: df). Odontophore ~ */s of buccal mass volume (Fig. 12: od). Odontophore muscles (Figs. 10-12, 16-20): mj, jaw and peribuccal muscle orig- inating on outer-ventral surface of odontophore cartilages (Fig. 20: mj), running towards dorsal region, splaying on dorsal wall of oral tube (Figs. 16-17, 19: my); ml, jugal muscles entirely covering haemocoelic structures, more concentrated close to the mouth; mlv, small pair of ven- tral retractor jugal muscles originating on lateral surface of haemocoel, close to the mouth, running towards medial region, inserting in ventral region of oral tube, close to the median line (Figs. 11, 12: mlv); mll, small pair of lateral retractor jugal muscles originating on dorso-lateral region of haemocoel, on the same level as oral tube, running dorso-ventrally, inserting in latero-dorsal surface of oral tube (Figs. 10, 11: m11); m2, strong pair of retractor mus- cles of buccal mass (or radular muscles) originating as one
Figs. 26-27. Leiostracus carnavalescus n. sp., anatomy. — 26. Central nervous system (nerve ring), in ventral view. 27. Central nerv- ous system (nerve ring), in dorsal view. — Scale bar: 1 mm.
pair of bundles of columellar muscle (Fig. 14: cm), running close to median line anteriorly along ~ 80 % of haemocoel length, inserting as two different bundles in ventropos- terior edge of odontophore, surrounding radular nucleus (Figs. 10, 16-17, 19-20: m2), passing through nerve ring (Fig. 10: nr); m3, transverse muscle of the posterior region of odontophore connecting both laterodorsal sides of odontophore, passing through odontophore posterior sur- face, in front of radular nucleus (Figs. 11, 16-17, 19: m3); m4, main pair of very thick dorsal tensor muscles of rad- ula originating on posteromedial region of odontophore cartilages, in 3-4 oblique layers (Figs. 17-18: m4), sur- rounding the cartilages, inserting in subradular membrane (Fig. 17: ir); m5, pair of thick auxiliary dorsal tensor mus- cles originating on posteroventral region of odontophore cartilages (Fig. 18:m5), along ~ 10% of their length, run- ning towards median line, inserting in subradular mem- brane by side of m4 insertion; m6, horizontal muscle thin, located between both cartilages in region just posterior to their fusion, along ~7/s cartilages length, keeping free ~ "/s of cartilages length posterior to it (Figs. 17-18, 20: m6); m7, small pair of muscles originating in middle level of cartilages central edge, just posterior to m6 (Fig. 20: m7),
18 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
running medial-posteriorly inside radular sac, inserting splaying in dorsal posterior inner surface of radular sac (Fig. 17:m7), m7a, small pair of muscles of similar char- acter as m7, but originating in posterior-medial corner of radular cartilages (Figs. 18, 20: m7a), running ventrally; m7b, small pair of narrow muscles originating in middle level of ventral edge of odontophore cartilages, covering m6, running posteriorly close to median line, inserting in dorsal inner surface of radular sac (Figs. 17-18: m7b); m11, pair of wide ventral tensor muscles of radula originating on ventrolateral surface of posterior region of cartilages (Fig. 20:m11), running close to the cartilages towards anterior region, inserting in ventral end of subradular membrane (br) (Fig. 17:m11); m13, pair of strong ventral odontophore protractor muscles originating on anteroven- tral region of haemocoel, running briefly towards dorsal region (Figs. 11, 12: m13), penetrating odontophore struc- tures by lateral aorta insertion, inserting in outer-pos- terior surface of odontophore cartilages (Fig. 20: m13). Odontophore non-muscular structures (Figs. 17—20): oc, odontophore cartilages flattened, elliptical, circa three times longer than wide, fused with each other for ~'/ of their anterior end (Figs. 18, 20: oc); anterior end roughly rounded; anterior region slightly curved medially; sc, sub- radular cartilage, with expanding region in buccal cavity protecting subradular membrane (Fig. 19: sc). Radular sac short, slightly extending beyond odontophore (Figs. 17, 19: rs).
Radula (Figs. 2-5) ~50% longer than odontophore; with rachidian teeth, and ~ 30 pairs of lateral teeth; no clear distinction between lateral and marginal teeth (Fig. 2). Rachidian tooth (Figs. 3-4: arrow) small, - '/o of radu- lar width and roughly twice longer than wide; base wider, barely triangular; cutting edge slightly rounded, with ~ 80 % of base length and 95 % of base width. Lateral teeth similar to rachidian (Figs. 3—4), except in being slightly asymmetrical, weakly arched towards median region, and with cutting edge circa twice the size of rachidian’s. Lat- eral teeth gradually decreasing towards lateral region; third or fourth teeth with gradually appearing basal-lateral cusp that becomes more and more distinct in more mar- ginal teeth. Marginal teeth starts with no clear boundary with lateral teeth; shaped similarly to lateral teeth, except for being smaller and with a proportionally larger second- ary cusp. There are more marginal teeth showing a bifid basal cusp (Fig. 5). Each radular row is disposed linearly on the same level (Fig. 2).
Salivary glands covering middle third of esophagus (Fig. 14: sg), forming two elongated white thin masses. Each salivary duct differentiable on anterior region of glands, with about 5-8% esophageal width. Salivary duct running on both sides of esophageal origin (Figs. 10, 14: sd), penetrating buccal mass wall on the region close to buccal ganglia (Fig. 16: sd). Salivary ducts opening on
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anterior middle level of dorsal folds, slightly separated from median line (Fig. 16: sa).
Esophagus one whorl long, with thin flaccid walls lacking clear subdivisions (Figs. 10—11, 14: es); inner sur- face simple, smooth. Posterior esophageal half slightly narrower than anterior half. Stomach (Fig. 14: st) rela- tively narrow, curved, somewhat fusiform: position and size described above (visceral mass); gastric walls thin, flaccid; inner surface smooth (Fig. 15), except for nar- row longitudinal fold (gf) from small gastric concavity up to intestinal origin, disappearing gradually. Esophageal insertion on right side, marked by sudden increase in size; intestinal origin on left side; both close to columella. Duct to anterior lobe of digestive gland encased between eso- phagus and intestine (Fig. 14: dd). Duct to posterior lobe of digestive gland located a short distance from intestinal or1- gin and posteriorly to the above-described duct, directed towards the opposite side (Fig. 14: dd). Intestine as wide as esophageal insertion along its entire length, flanking the left side of the anterior lobe of digestive gland; on the region of the kidney, it gradually turns right and anteri- orly, running almost straightly forward in the pallial cav- ity (Fig. 14: in). Rectum and anus described above (pallial cavity; Fig. 7).
Genital system (Figs. 21-25): gonad described above (visceral mass). Hermaphroditic duct (Fig. 11: hd) narrow and weakly coiled; running for ~ whorl close to columella. Seminal receptacle (Figs. 22—23: sr) elongated, sac-lıke, strongly curved, with circa three times the hermaphroditic duct width, and ~10% its length. Fertilization complex simple, located on the base of seminal receptacle (Fig. 23) as a continuation of the hermaphroditic duct and the sem- inal receptacle’s duct; ~ 2 width and length of receptacle. Fertilization complex totally immersed in albumen gland (Fig. 22), inserting in the posterior end of spermoviduct, in the albumen gland duct’s region; all of them of similar width. Albumen gland (Figs. 21—23: ag) solid, white, ellip- tical, as large as gonad (~'/s whorl). Albumen gland duct subterminal, connected to distal end of spermoviduct, pre- ceded by pair of large albumen chambers (Figs. 21—23: ac), each with ~ 4 albumen gland size, located on dorsal region of transition between albumen gland and spermoviduct, successively connected to the beginning of spermoviduct (Fig. 23). Spermoviduct ~ 1.5 whorl in length, slightly nar- rower than albumen gland, ~20 times longer than wide. Prostate gland occupying ~ % of the spermoviduct volume (Fig. 23: pt). Uterus occupying the other half of spermo- viduct volume; external walls thick-glandular (Fig. 23: ut); inner surface completely covered by ample transversal folds (Fig. 25: ut); posterior diverticulum well-developed on the region immediately anterior to the albumen cham- bers (Fig. 23: gd). Sperm grove simple in posterior two thirds of spermoviduct (Fig. 13: sp), protected ventrally by a tall fold; gradually becoming tubular fold (Fig. 25: fd) as
SIMONE & SALVADOR, TAXONOMICAL STUDY ON A SAMPLE OF LAND SNAILS FROM NANUQUE 19
vas deferens, along anterior third of spermoviduct, with ~'/s of anterior spermoviduct width; bearing clear swell- ing (Fig. 25: vf) on region preceding exteriorization of vas deferens (Figs. 21, 25: vd). Vagina ~'/io of spermo- viduct length; inner surface simple, with 4-5 longitudi- nal, low, wide folds (Fig. 25: vg). Bursa copulatrix as long as spermoviduct plus albumen gland; bursa duct as wide as adjacent spermoviduct on its origin but gradually nar- rowing towards posterior end (Figs.21, 25: bd); bursa elliptical, -% of albumen gland size (Fig. 21: bc), located encased between pericardium and adjacent intestinal loop. Penis ~4/s of spermoviduct length, -% its anterior width (Fig. 21: pe); penis muscle inserting terminally, very short (Fig. 21: pm). No epiphallus. Vas deferens inserted sub- terminally in penis tip (Figs. 21, 24: vd). Internal penial surface with 3 clear sub-chambers (Fig. 24): posterior sub- chamber with ~ penis length, bearing 4—5 longitudinal narrow low well-separated folds, some of them converg- ing to the vas deferens aperture (vd), anterior quarter with fold becoming oblique and interrupted (pf); middle sub- chamber (p1) separated from other sub-chambers by trans- verse constrictions (posterior constriction slightly taller than anterior one), with ~ % penis length, internal surface bearing 10-12 longitudinal narrow closely-packed folds; anterior sub-chamber narrow, with ~'% of penis length, bearing 10-12 inner longitudinal low closely-packed folds. Penis shield with same length as anterior penial sub- chamber (Fig. 24: ps). Genital pore round, simple.
Central nervous system (Figs. 26-27): located dor- sally from ventral base of buccal mass up to the transition between buccal mass and esophagus (Figs. 10, 14: nr). Pair of cerebral ganglia (ce) largely fused with each other; cere- bral commissure invisible; each ganglion about as wide as adjacent esophageal section; several wide nerves origi- nating on cerebral anterior edge, including relatively nar- row subesophageal ganglion (su) located close to cerebral ganglia (but not fused with it). Pair of optical ganglia not individualized, possibly part of other nerves. Four paral- lel connectives (cn) between cerebral ganglia and pedal ganglia. Pair of pedal ganglia (pp) forming a single mass located opposite to the cerebral ganglia, slightly smaller in size than the latter. No differentiable ganglion detectable except for medial weak constriction. Several pairs of pedal nerves originating from latero-anterior corner of these ganglia. Anterior aorta (aa) clearly inserted ın anteroven- tral region between both cerebral ganglia. Pair of stato- cysts not seen.
Measurements (in mm). Holotype: 8 whorls; H=25.4; D=12.4; S=16.9; S’ =13.4; h=9.2; d=8.1. Mean (n=17): 7 to 8 whorls; H=244+1.3 (max 26.6; min 22.8); D=I3.L409 “max: 16.0 min 12:1): S$ =14.92 0:9 “(max 16.9; min 13.6), S’=11.7+0.9 (max 13.4; min 10.4); h=10.0+0.8 (max 12.1; min 8.9); d=7.6+0.4 (max 8.3; min 6.9).
Distribution
Brazil. Minas Gerais: Nanuque (type locality) and Mantena municipalities. Espirito Santo: Pinheiros and Sooretama municipalities.
Habitat
Found on trees or tall bushes in Nanuque. Specimens from Mantena were found on subshrubs; specimens from Espirito Santo state have no habitat data. It is unknown whether its remarkable color pattern is related to camou- flage, aposematism, or if it indeed plays some role on the animal’s life at all. Still, ıt should be noticed that many specimens show signs of predation by birds.
Remarks
Leiostracus carnavalescus n.sp. bears some resem- blance to L. obliquus (Reeve, 1849), from Bahia and Minas Gerais states (SIMONE 2006); however, it has a much thinner shell, a much less reflected peristome, wider and more conical shell, and the remarkable and variable color pattern (see description above). Shells with variable color pattern are not uncommon in the genus, but usually com- prise only varying tones of yellow and brown (e. g., SIMONE 2006, SALVADOR & CAVALLARI 2013). The species which more closely resembles the pattern of L. carnavalescus is L. vittatus (Spix, 1827): it is white with one to three spiral brownish bands and some specimens show an “inverted” color pattern (base color brown with white spiral bands); moreover, it occurs in the states of Pernambuco and Bahia (SIMONE 2006), a neighboring distribution in relation to L. carnavalescus. Still, L. carnavalescus can be easily dis- tinguished by its wider and more conical shell, with a smaller and rounder aperture.
The specimens from the municipality of Mantena (Fig. 1), approximately 130km to the south of Nanuque, seem to belong to the same species as far as the concho- logical characters indicate. However, slight differences in coloration are evident (Figs. 64—65), although their impor- tance cannot be evaluated without additional material. The specimens from Mantena are white with only two spi- ral bands on the body whorl (one red and the other, close to the umbilicus, dark brown), have an yellowish brown spire top and have only a single line of yellowish brown dots on their spire (these dots can be very large in some instances: Fig. 64). Moreover, in one specimen (Fig. 64), the suture line is marked with a brown line. Finally, while the ani- mals from Nanuque were found on trees or tall bushes, the ones from Mantena were supposedly found on subshrubs, at least according to the specimens’ labels.
The specimens from Espirito Santo state (Figs. 62—63), on the other hand, present a color pattern extremely simi- lar to the type material from Nanuque; the single exception is that these specimens lack the spiral line of dots, having
20 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
a solid line instead. The municipality of Pinheiros is circa 65 km to the southeast of Nanuque, while the municipal- ity of Sooretama is circa 150km to the southeast (Fig. 1). Unfortunately, these specimens have no habitat data.
Anatomically, LZ. carnavalescus has the normal pat- tern of the Bulimulinae (BREURE & SCHOUTEN 1985; SIMONE 1998). The more remarkable features are the multiplicity of the columellar muscle bundles (Fig. 14: cm); the enlarge- ment of the pedal gland (Fig. 14: pg); the multiplicity of the pair of odontophore muscles m7 (inside radular sac: m7, m7a, m7b; Figs. 17-18, 20); the pair of ventral protractor muscles of odontophore (Figs. 11, 12: m13); the two albu- men chambers (Fig. 22: ac); the simplicity of the fertilization chamber (Fig. 23); and the penis divided into 3 sub-cham- bers (Fig. 24). The significance of these differences, in spe- cies, genus and family levels, is still under analysis.
Genus Rhinus Martens in Albers, 1860
Rhinus botocudus n. sp. (Figs. 28-52, 68-76)
Type material
Holotype: MZSP 106174 (Figs. 68-73). Paratypes: MZSP 106175 (2 specimens), 106176 (10 shells), from type locality. J. Corrro col., May/2012.
Type locality: Brazil. Minas Gerais; Nanuque, ~ 120 m of elevation (~ 17°51'S 40°23'W).
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Etymology
Reference to the “Botocudos” native Indians who once dwelt in the region.
Diagnosis Shell small and globose; aperture large and elongated diagonally. Protoconch with different sculpture patterns on upper and lateral portions of the whorl. Whitish spiral line running on the middle portion of body whorl.
Description
Shell (Figs. 68—74): medium-sized, ovoid to globose; diameter -% shell length. Color yellowish brown to light brown, with whitish spiral line on middle portion of body whorl (Fig. 68). Spire angle ~ 75°. Protoconch of 1.5 whorl, sculptured by axial parallel riblets on upper portion of whorl and spiral parallel striae on lateral portion of whorl (Fig. 72); transition to teleoconch clear, orthocline. Teleo- conch smooth, except for growth lines, with lightly hairy periostracum; hairs arranged in spiral parallel lines, ~ 45 on body whorl, uniformly spaced and distributed (Fig. 73). Whorl profile convex. Suture well-marked. Aperture large, oval, diagonally elongated, slightly prosocline; ~?/s shell length, -% shell width. Peristome slightly reflexed, more reflexed on columellar region, partially covering umbili- cus. Body whorl -% shell length. Umbilicus narrow.
Head-foot (Figs. 37, 38, 74): features similar to Leio- stracus carnavalescus (see above), with the following
Figs. 28-31. Rhinus botocudus n.sp., SEM images of radulae. The arrows indicate the rachidian column. — 28. Whole view. 29. Detail of central region. 30. Greater detail of central region. 31. Detail of marginal region. — Scale bars: 20 um (30, 31), 50 um (29), 200 um (28).
SIMONE & SALVADOR, TAXONOMICAL STUDY ON A SAMPLE OF LAND SNAILS FROM NANUQUE 21
differences: Color uniformly dark grey on exposed areas (Fig. 74). Pair of ommatophores retractor muscles elon- gated, originating from columellar muscle (Fig. 37: mu). Columellar muscle of 1.5 whorl in length, divided into 4—5 separated bundles (Figs. 37-38: cm).
Mantle organs (Figs. 32-35): features similar to Leio- stracus carnavalescus (see above), with the following dif- ferences: Mantle border thick and very tall (Fig. 32: mb). Secondary fold of mantle border located on bottom side of mantle collar (Figs. 32-33: mf). Outer pneumostome fold with circa twice the length of inner fold. Anus is a sep- arated aperture located to the right of the pneumostome (Figs. 32, 34: an). Lung relatively wide. Anterior three quarters of pulmonary vessels conspicuous (Fig. 32); pos- terior quarter nearly smooth; venation absent on the region close to pneumostome (Fig. 34). Kidney occupying ~ 30 % of cavity length and ~ 50 % of its width.
Visceral mass (Fig. 39): features similar to Leiostracus carnavalescus (see above), with the following differences: Visceral mass ~ 2.5 whorls in length; stomach ~ '/s of vis- ceral volume (Fig. 39: st).
Circulatory and excretory systems (Figs. 32-35): fea- tures similar to Leiostracus carnavalescus (see above), with the following differences: Pericardium occupying ~ 10% of lung area. Kidney with coiled kidney lobe (“rou- lade-like”; Fig. 36: kl). Nephropore punctiform, located on anterior-left corner (Fig. 35: ne). Urinary aperture wide (Fig. 32: ua). Urinary gutter well-delimited by pair of folds, running parallel to mantel border (Fig. 34: ug).
Digestive system (Figs. 38-45): features similar to Leiostracus carnavalescus (see above), with the follow- ing differences: Jaw with constant width along its length, bearing fewer transverse folds; median folds more perpen- dicular to edges (Figs. 40: jw, 75-76); color yellow, trans- lucent. Buccal mass occupying ~'/s of haemocoel volume (Fig. 38: bm). Pair of dorsal folds or oral cavity slightly narrower (Fig. 40: df). Salivary apertures (sa) located slightly more posteriorly. Differences in odontophore mus- cles (Figs. 40—45): mj, origin more elongated and posteri- orly located (Fig. 45: mj); mlv and mll pairs undetectable; mld, pair of small dorsal protractor muscles (Fig. 41: mld) originating on dorsal peribuccal inner surface of haemo- coel, running to posterior region flanking buccal mass and inserting in posterior-dorsal surface of buccal mass; m2, slightly narrower near insertion (Figs. 42—43: m2), insert- ing in m4 instead of cartilages; m3, narrower (Fig. 40: m3); m3p, pair of thin and narrow muscles with dorsoventral fibers connecting lateral region of esophageal origin with ventral region of m2 insertion (Figs. 40-41: m3p); m4, slightly thicker (Figs. 42—43: m4); m5, shorter and thinner (Fig. 43: m5), m6, narrower on posterior region (Figs. 43, 45: m6); m7, m7a and m7b undetectable; m11, originating more anteriorly, narrower near origin (Fig. 45: m11); m13, slightly thicker near origin (Fig. 45: m13), but quickly nar-
rowing (Figs. 41—42:m13); mx, strong thick muscular layer produced medially by mj pair, passing posteriorly to both odontophore cartilages (Fig. 42: mx). Differences in odontophore’s non-muscular structures (Figs. 42—45): odontophore cartilages’ anterior junction slightly more pointed (Fig. 43: oc), fused with each other along ~'/s of their length, on their anterior end (Figs. 44—45: of).
Differences in radula (Figs. 28-31): ~40 pairs of lat- eral teeth; distinction between lateral and marginal teeth somewhat clear, marked by abrupt inclination of tooth row (Fig. 28). Rachidian tooth (Figs. 29-30: arrow) ~ !/so of radular width and circa twice longer than wide; base with circa twice the length of cutting edge; cutting edge somewhat triangular, pointed. Lateral teeth (Figs. 29-30) always with secondary lateral cusp (of ~’2 main medial cusp size); ~10 pairs of lateral teeth per row. Marginal teeth with 2—3 small secondary cusps on main cusp; basal cusp small, slightly bifid (Fig. 31).
Esophagus lacking clear subdivisions along its length (Fig. 39: es); lacking duct to anterior lobe of digestive gland. Stomach (Fig. 39: st) spherical; gastric walls thin; inner surface bearing uniform cover of narrow longitudi- nal folds. Duct to anterior lobe of digestive gland encased between esophagus and intestine, opening directly into anterior region of stomach (Fig. 39: dd). Duct to posterior region located on the middle of posterior gastric surface.
Genital system (Figs. 46-50): features similar to Leio- stracus carnavalescus (see above), with the following dif- ferences: Gonad with more distinct lobes (Fig. 46: go). Hermaphroditic duct shorter, less coiled and much thicker (Figs. 38, 46: hd). Insertion of hermaphroditic duct very narrow (Fig. 47: hd). Seminal receptacle narrower, coiled, ~10times longer than wide (Fig. 47: sr). Fertilization complex about as wide as seminal receptacle and ~ % its length; anterior end of complex with wide albumen gland duct and pair of albumen chamber ducts on opposite side (Fig. 47: ad, ac). Albumen chambers (Figs. 46-47: ac) each of ~'/s albumen gland size; the most posteriorly located chamber 1s duplicated. Spermoviduct slightly shorter and broader, of ~1 whorl in length, ~12times longer than wide. Prostate gland occupying ~ 4 of spermoviduct vol- ume (Fig. 46: pt). Uterus occupying other % of spermo- viduct space; external walls with strong transverse folds (Fig. 46: ut); no posterior diverticulum (Fig. 47). Sperm grove is a tall fold on almost the entirety its length (Figs. 46, 48: fd); vas deferens exteriorization simple, con- nected to internal fold (Fig. 48: vf). Bursa copulatrix as long as spermoviduct; bursa duct as wide as adjacent sper- moviduct near its origin, but gradually narrowing towards posterior end (Figs. 46, 48: bd), up to ~'/s spermoviduct width; bursa with ~'/s of albumen gland size (Fig. 46: bc); one specimen with bulging due to fusiform, elongated spermatophore (Fig.46:bd). Penis - spermoviduct length (Figs. 38, 46: pe); penis muscle short, inserting
22 STUTTGARTER BEITRÄGE ZUR NATURKUNDE A Neue Serie 9
Figs. 32-38. Rhinus botocudus n. sp., anatomy. 32. Pallial cavity, in inner ventral view (ventral flap of pneumostome [pv] deflected). 33. Mantle border, in frontal view (as in situ). 34. Detail of pneumostome region, in ventral view (ventral flap of pneumostome [pv] deflected upwards). 35. Posterior region of pallial cavity, in inner ventral view (heart and reno-pericardial apertures seen by transpar- ency). 36. Transverse section of the middle level of kidney (ventral side to the right). 37. Head-foot, in dorsal view (most of the dorsal side of haemocoel removed to show columellar muscle components). 38. Head-foot and visceral mass, mostly in ventral view (dorsal wall of haemocoel removed; pallial cavity partially shown; most structures seen as in situ). — Scale bars: 2mm.
SIMONE & SALVADOR, TAXONOMICAL STUDY ON A SAMPLE OF LAND SNAILS FROM NANUQUE
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Figs. 39-45. Rhinus botocudus n. sp., anatomy. — 39. Foregut and midgut, in dorsal view (some adjacent structures are also shown). 40. Buccal mass, in ventral view (odontophore partially removed and deflected to the right). 41. Buccal mass, in lateral (right) view. 42. Odontophore, in ventral view (right intrinsic muscles deflected; part of subradular membrane [br] removed on the right side).
43. Odontophore, in dorsal view (radula still partially connected, but deflected downwards). 44. Right odontophore cartilage, in dor- sal view, showing origin of the intrinsic muscles. 45. Right odontophore cartilage, in ventral view. — Scale bars: 2mm.
23
24 STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
Figs. 46-52. Rhinus botocudus n. sp., anatomy. — 46. Genital system, in ventral view (with cross-section of indicated portion of sper- moviduct). 47. Genital system, detail of region of carrefour (albumen gland [ag] seen as transparent structure). 48. Terminal region of genital system, in dorsal view (most of the female portion opened longitudinally). 49. Penis, detail of distal region. 50. Penis, in dor- sal view, entirely opened longitudinally. 51. Central nervous system (nerve ring), in ventral view. 52. Central nervous system (nerve ring), in dorsal view. — Scale bars: 1 mm.
SIMONE & SALVADOR, TAXONOMICAL STUDY ON A SAMPLE OF LAND SNAILS FROM NANUQUE 25
subterminally (Figs. 49-50: pm). Epiphallus is a simple blind sac, ~ % penis length (Figs. 49-50: eh). Vas deferens inserted between epiphallus and penis (Figs. 49-50: vd). Internal penial surface lacking clear inner divisions, except for anterior wide short region (Fig. 50: pi); 5-6 lon- gitudinal simple wide folds, closely packed together, but gradually becoming narrower and more distanced from each other towards anterior chamber (Fig. 50). Penis shield absent.
Central nervous system (Figs. 51, 52): features sim- ilar to Leiostracus carnavalescus (see above), with the following differences: Central nervous system located more anteriorly, mostly on the ventral base of buccal mass (Fig. 39: nr). Ganglia more separated and more individu- alized. Pair of cerebral ganglia (ce) more separated from each other; cerebral commissure about as long as each gan- glion; each ganglion with about half the size of adjacent esophageal section. Between cerebral ganglia and pedal ganglia, there are three parallel symmetrical connectives (cn), each circa three times as long as cerebral ganglia. Pair of pedal ganglia (pp) forming single mass, about as large as cerebral ganglia; weak constriction between the two pedal ganglia. Statocysts with several iridescent inter- nal granules (Fig. 51: cy). Buccal ganglia well-developed, located laterally between esophageal origin and odonto- phore (Fig. 40: bg).
Measurements (in mm). Holotype: 4.75 whorls; H=13.9; D=11.7; S=5.6, S’=3.4; h=8.7; d=7.6. Mean (n=9): ~4.5 whorls; H=14.7+0.8 (max 15.9; min 13.9); D=11.5+0.6 (max 12.3; min 10.4); S=6.0+0.3 (max 6.4; min 5.4); S’=3.5+0.3 (max 3.8; min 2.9); h=9.2+0.5 (max 10.1; min 8.4); d=7.4+0.5 (max 8.2; min 6.6).
Distribution Known only from type locality.
Habitat Found on low foliage.
Remarks
Rhinus botocudus n.sp. is reminiscent of only two Brazilian species: R. velutinohispidus (Moricand, 1836) and R. /ongisetus (Moricand, 1846), both from Bahia state (SIMONE 2006). Rhinus botocudus can be distinguished from R. velutinohispidus by being much smaller, barely reaching half the size of that species, and having a deeper suture and more convex whorls. Rhinus botocudus can be distinguished from R. /ongisetus by being larger, having a narrower spire and a more laterally positioned body whorl. Finally, R. botocudus has an arrangement of hairs on the periostracum similar to R. ciliatus (Gould, 1846), from Rio de Janeiro state (SIMONE 2006); nevertheless, it can be differentiated from this species by its smaller size, more rounded shell profile, lower spire, a whitish band on the
middle portion of the body whorl and more separate spi- ral lines of hair.
Rhinus ıs an endemic South American genus, occur- ring in Venezuela and Brazil, being particularly diverse in the latter (BREURE 1979, SımonE 2006). Anatomi- cally, the distinctive description above explored the main exclusivities of R. botocudus when compared with the known orthalicoids, and with Leiostracus carnavalescus described above. Particularly, the more remarkable exclu- sivities are: the origin of the tentacular muscle ın the colu- mellar muscle (Fig. 37); the absence of the esophageal duct to the anterior lobe of the digestive gland, which instead is located in the stomach (Fig. 39); the presence of three albumen chambers (Fig. 46); and the simple internal sur- face of the penis (Fig. 50). Additionally, of great interest are the multiple columellar muscles (Fig. 37: cm), similar to what is seen in Leiostracus carnavalescus. The analy- sis of these differences and shared features is still a matter of ongoing analysis.
Family Subulinidae Genus Obeliscus Beck, 1837
Obeliscus boitata n. sp. (Figs. 77-79)
Type material
Holotype: MZSP 106169 (Figs. 77-79). Paraty pes: MZSP 106170 (39 shells), from type locality. J. Cottro col., May/2012.
Type locality: Brazil. Minas Gerais: Nanuque, ~ 120 m of elevation (- 17°51'S 40°23'W).
Etymology
The name (in apposition) is a reference to Boitata, a fiery snake-like creature that protected the forests in Brazilian folklore.
Distribution Known only from type locality.
Diagnosis Spire top narrow, elongated. Spire apex (protoconch) bulbous, domed, thicker than subsequent spire portion. Suture perpendicular to shell length axis. Whorls with a faint keel on median region. Aperture rectangular.
Description
Shell large, multispiral, thin, conical, imperforate; Spire top narrow, elongated. Spire apex (protoconch) bulbous, domed, thicker than subsequent spire por- tion; diameter ~'%4 shell length. Color translucent white to dirty white. Spire angle ~15°. Protoconch of -2.75 whorls, rounded, dome-shaped, blunt, smooth; transition
26 STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
Figs. 53-67. Leiostracus carnavalescus n.sp. — 53-57. Holotype (MZSP 106177; H=25.4mm). 58. Holotype, protoconch detail (inferior side = 2mm). 59. Paratype #1 (MZSP 106179). 60. Paratype #2 (MZSP 106178; H=21.2mm). 61. Paratype #3 (MZSP 106179). 62. Specimen #1 from Pinheiros, Espirito Santo (MZSP 106618; H=21.2 mm). 63. Specimen #2 (juvenile) from Sooretama, Espirito Santo (MZSP 106666; H= 10.3 mm). 64. Specimen #1 from Mantena, Minas Gerais (MZSP 108010). 65. Specimen #2 from Mantena, Minas Gerais (MZSP 108010). 66. Detail of mouth showing exposed portion of jaw (inferior side = 3 mm). 67. Living spec- imen, paratype #4 (MZSP 106178).
SIMONE & SALVADOR, TAXONOMICAL STUDY ON A SAMPLE OF LAND SNAILS FROM NANUQUE DH
Figs. 68-76. Rhinus botocudus n. sp. — 68-71. Holotype (MZSP 106174; H= 13.9 mm, D= 11.7 mm). 72. Same, detail of shell apex (inferior side =2 mm). 73. Same, detail of surface in middle-right side of last whorl, showing hairy periostracum (right side = 3 mm). 74. Living specimen, paratype (MZSP 106175, H= 15.3 mm). 75. Extracted jaw, in anterior to slightly posterior view (width = 1.2 mm). 76. Jaw, as seen in situ, in anterior view.
28 STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
Figs. 77-100. Land snails from Nanuque, Minas Gerais State, Brazil. — 77-79. Obeliscus boitata n. sp., holotype (MZSP 106169; H=50.3 mm). 80. Auris bilabiata (MZSP 106155; H=42 mm). 81-83. Helicina boettgeri (MZSP 106166; D=4.1 mm). 84-86. Heli- cina variabilis, specimen #1 (MZSP 106171; D=12.9mm). 87-89. Helicina variabilis, specimen #2 (MZSP 106172; D= 12.5 mm). 90-92. Cyclopomops moricandi (MZSP 106168; D=5.1 mm). 93-95. Rectartemon piquetensis (MZSP 106156; D=31.6 mm). 96. Bahiensis cf. bahiensis (MZSP 106165; H=16.2 mm). 97. Dysopeas muibum (MZSP 106163; H=5.7 mm). 98-100. Prohappia besckei (MZSP 106164; D=7.1 mm).
SIMONE & SALVADOR, TAXONOMICAL STUDY ON A SAMPLE OF LAND SNAILS FROM NANUQUE 29
to teleoconch clear. Teleoconch sculptured by numerous thin axial striae, stronger on the upper portion of whorl near the suture. Whorl profile slightly convex; faint keel on median region. Suture well-marked. Aperture small, prosocline, oval to rectangular; ~!/s shell length, ~ 2 shell width. Peristome simple, weakly reflexed. Body whorl ~ YA Shell length.
Measurements (in mm). Holotype: 13.75 whorls; H=50.3, D=11.9; S=40.9; S’ =35.9; h=9.7; d=6.1. Mean (n= 10): from 13.5 to 14.5 whorls; H=49.1 + 1.9 (max 52.0; min 47.0); D=11.5+0.4 (max 11.9; min 10.8); S=40.1+1.8 (max 43.2; min 38.0), S’ =35.6+ 1.3 (max 37.9; min 33.7); h=94+0.4 (max 10.0; min 8.6), d=6.0+0.3 (max 6.4; min 5.6).
Remarks
Obeliscus boitata n.sp. closely resembles, in overall Shell shape and color, only O. carphodes (Pfeiffer, 1852) and O. sylvaticus (Wagner, 1827), the known distribu- tion of the latter 1s given as a vague “northeastern Bra- zil”, while the former’s is even vaguer, given simply as “Brazil” (SIMONE 2006). Obeliscus boitata n.sp. can be distinguished from both these species by its narrow and elongated spire, bulbous protoconch, faint keel and a suture more perpendicular to the shell length’s axis. It can be further differentiated from O. carphodes by its smaller size and rectangular aperture and from O. sy/vaticus by its larger size and shorter whorls.
4 Discussion
Besides the new species described above, a list of the other species found in the land snail sample from Nanuque can be seen on Tab. 1. The records of some species pre- sented here are the first ones for Minas Gerais state: Auris bilabiata (Broderip & Sowerby, 1829) (Fig. 80), Helicina boettgeri Wagner, 1910 (Figs. 81-83), Helicina variabi- lis Wagner, 1827 (Figs. 84—89), Cyclopomops moricandi (Pfeiffer, 1852) (Figs. 90-92), Rectartemon piqueten- sis (Pilsbry, 1930) (Figs. 93-95), Bahiensis cf. bahien- sis (Moricand, 1833) (Fig. 96), Dysopeas muibum Marcus & Marcus, 1968 (Fig.97) and Prohappia besckei (Dun- ker in Pfeiffer, 1847) (Figs. 98-100). Although Nanuque is located very close to the borders with Bahia and Espirito Sato states (Fig. 1), the present records still represent an extension in the distribution of these species (Tab. 1). Most of the species presented here belong to genera already well recorded for the Brazilian southeast region, but three gen- era in particular (Cyclopomops Bartsch & Morrison, 1942, Dysopeas Baker, 1927 and Prohappia Thiele, 1927) are completely new records for Minas Gerais.
The natural environment around Nanuque was severely degraded by incessant woodcutting, crops, breeding of livestock and the construction of railroads
and highways since the final decades of the 19" century (CERQUEIRA NETO 2005, Martins 2010). Nevertheless, the single forest fragment studied here still harbors a unique fauna and even new species. Our findings in Nanuque are a reminder of how these remnants of Atlantic rainforest might be acting as refuges (e. g., PEREIRA et al. 2014), har- boring still unknown diversity and perhaps even their own share of endemic species. Despite the regions’ flora and fauna being scarcely studied, a high level of biodiversity is thought to be the norm for the broader region encom- passing Nanuque and its surroundings and concerns about extinction have already been raised for plant and fish spe- cies (CERQUEIRA NETO 2005, SARMENTO-SOARES et al. 2007).
Alarmingly, José CoLrro Jr. and his team returned much more recently to the same forest fragment in Nanuque, only to discover that it was nearly entirely devastated in order to open space for more tomato crops. Whether the new species described here have already become extinct remains to be seen, as they may still live unknown and undisturbed in other localities (Leiostracus carnavales- cus, for instance, also occurs elsewhere). Still, ıt 1s well- known that many species are becoming extinct before been even known to science (e. g., HAWKSWORTH & CoWIE 2013), even in groups prone to greater study and protection efforts, like birds (e. g., PEREIRA et al. 2014). Therefore, the situation might be much direr for land and freshwa- ter mollusks, which are deemed the most imperiled group of animals, with the highest extinction rates (LYDEARD et al. 2004, REGNIER et al. 2008). Despite basic science being currently out of vogue, a more thorough knowledge of the local fauna and flora of these critical places ıs the first step towards protection and appropriate legislation.
5 References
BREURE, A. S. H. (1979): Systematics, phylogeny and zoogeogra- phy of Bulimulinae (Mollusca). — Zoologische Verhandelin- gen 168: 1-215.
BREURE, A. S.H. & SCHOUTEN, J. R. (1985): Notes on and descrip- tions of Bulimulidae (Mollusca, Gastropoda), III. — Zoolo- gische Verhandelingen 216: 1-98.
CERQUEIRA NETO, S. P. G. (2005): Contribuicäo ao estudo geogra- fico do municipio de Nanuque (MG). — Caminhos de Geo- grafia 9(15): 82-92.
HAwKSWORTH, D.L. & CowiE, R.H. (2013): The discovery of historically extinct, but hitherto undescribed, species: an under-appreciated element in extinction-rate assessments. — Biodiversity and Conservation 22: 2429-2432.
LYDEARD, C., CowIE, R. H., PONDER, W. F., BoGAN, A. E., BOUCHET, P., CLARK, S.A., CumMInGs, K.S., FREST, T. J., GARGOMINY, O., HERBERT, D.G., HERSHLER, R., PEREZ, K.E., RoTH, B., SEDDON, M., STRONG, E.E. & THompson, F. G. (2004): The Global Decline of Nonmarine Mollusks. — BioScience 54: 321-330.
Martins, M.L. (2010): Ocupagäo e desflorestamento numa area de fronteira: Vale do Mucuri, MG: 1890 a 1950. — Revista de Histöria Regional 15: 40-77.
30 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
MORRETES, F. L. (1949): Ensaio de catälogo dos moluscos do Bra- sil. — Arquivos do Museu Paranaense 7: 5-216.
PEREIRA, G.A., Dantas, S.M., SILVEIRA, L.F., Ropa, S.A., ALBANO, C., SONNTAG, F.A., LEAL, S., PERIQUITOo, M.C., Ma acco, G. B. & Lees, A. C. (2014): Status of the globally threatened forest birds of northeast Brazil. — Papéis Avulsos de Zoologia 54 (14): 177-194.
REGNIER, C., FONTAINE, B. & Boucuet, P. (2008): Not know- ing, not recording, not listing: numerous unnoticed mollusk extinctions. — Conservation Biology 23: 1214-1221.
Rosinson, D. G. & SLAPCINSKY, J. (2005): Recent introductions of alien land snails into North America. — American Malaco- logical Bulletin 20: 89-93.
Authors’ addresses:
Neue Serie 9
SALVADOR, R. B. & CAVALLARI, D.C. (2013): Taxonomic revision of Leiostracus onager and Leiostracus subtuszonatus (Gas- tropoda: Pulmonata: Orthalicidae). — Journal of Conchology 41: 511-518.
SARMENTO-SOARES, L.M., Mazzont, R. & MARTINS-PINHEIRO, R.F. (2007): A fauna de peixes na bacia do Rio Peruipe, extremo Sul da Bahia. — Biota Neotropica 7: 291-308.
SIMONE, L.R.L. (1998): Anatomical description of Anctus angi- ostomus (Wagner, 1827) from northeastern Bahia, Brazil (Gastropoda, Pulmonata, Bulimulidae). — Studies on Neo- tropical Fauna and Environment 33: 170-177.
SIMONE, L.R.L. (2006): Land and freshwater mollusks of Brazil, 390 pp.; Sao Paulo (Editora Grafica Bernardi & Fundacäo de Amparo a Pesquisa do Estado de Säo Paulo).
Luiz RıcARDO Lopes SIMONE, Museu de Zoologia da Universidade de Säo Paulo, Avenida Nazare 481, 04218-970 Sao Paulo, Brazil;
e-mail: Irsimone@usp.br
RoDRIGO BRINCALEPE SALVADOR (corresponding author), Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191, Stutt- gart, Germany or Mathematisch-Naturwissenschaftliche Fakultät, Eberhard Karls Universität Tübingen, Hölderlinstraße 12, 72074
Tübingen, Germany; e-mail: salvador.rodrigo.b@gmail.com
Manuscript received: 22.1X.2015, accepted: 25.X1.2015.
Stuttgarter Beiträge zur Naturkunde A, Neue Serie 9: 31-39; Stuttgart, 30.1V.2016. DOI: 10.18476/sbna.v9.a3 31
Two new species and new records of terrestrial isopods (Isopoda: Oniscidea) from Oman
HELMUT SCHMALFUSS
Abstract
A small but highly interesting collection of terrestrial isopods from Oman is reported. The new species Xeronis- cus siegfriedhuberi n. sp. and Somalodilloides pseudopilosus n. sp. are described and illustrated, and new records are given for the species Chaetophiloscia sp., Periscyphis omanensis, Xeroniscus erythraeus and X. troglophilus.
Key words: Isopoda, Oniscidea, Oman, new species, new records.
Zusammenfassung
In einer kleinen, aber hochinteressanten Aufsammlung von Landisopoden aus dem Oman fanden sich die neuen Arten Xeroniscus siegfriedhuberi n.sp. und Somalodilloides pseudopilosus n.sp., die beschrieben und abgebil- det werden, und neue Nachweise von Chaetophiloscia sp., Periscyphis omanensis, Xeroniscus erythraeus and X.
troglophilus.
Contents ja Tntrodiekonen. ME Seren AR U BR Wan a EI FE SE UF Ae eA BE BER 2 EIER 31 DAN EOS Rn Bin ea te a a A a Me a a atte nn deem N in tat ta tet De ee 32 3, "Newiterrestrial 1s@ pod material KM Oman. Au na tle ae en Fee kennen rer 32 4, SRCTCRETIOCOS UL re, Se N ana In, eee Nee Auen Serie nee A nen Mn Me EM 39
1 Introduction
Until the year 1988 only one species of the Oniscidea was known from Oman (Tylos maindroni Giordani Soika, 1954). FERRARA & Tait (1988) reported six additional spe- cies from the country. In the most recent publication on Oman’s terrestrial isopods (Taiti et al. 2000) a number of additional species are recorded, adding up to 17 species known from the country.
In the year 2000 SIEGFRIED HUBER accompanied Prof. PETER WEYGOLDT on a collecting tour through Oman and collected a number of isopod samples. This new material contains two species new to science and new records of four additional species, two of which are reported for the first time from Oman, thus extending their distribution area considerably.
The new records increase the number of terrestrial iso- pod species known from this country to the following 21 species:
Ligia pigmentata Jackson, 1922
Tylos maindroni Giordano Soika, 1954 Chaetophiloscia sp.
Agnara gallagheri (Ferrara & Taiti, 1988) Agnara madagascariensis (Budde-Lund, 1885) Porcellionides pruinosus (Brandt, 1833)
Agabiformius lentus (Budde-Lund, 1885)
Proporcellio vulcanius Verhoeff, 1908 (= P. quadriseriatus) Porcellio evansi Omer-Cooper, 1923
Porcellio sp.
Periscyphis vittatus (Omer-Cooper, 1926)
Periscyphis albomarginatus Taiti, Ferrara & Allspach, 1997 Periscyphis insularis Ferrara & Taiti, 1988
Periscyphis omanensis Taiti & Ferrara, 1991
Periscyphis dhofarensis Taiti, Ferrara & Davolos, 2000 Xeroniscus troglophilus Taiti, Ferrara & Davolos, 2000 Xeroniscus erythraeus (Ferrara, 1972)
Xeroniscus sieg friedhuberi n. sp.
Omanodillo gardneri Taiti, Ferrara & Davolos, 2000 Somalodillo paeninsulae Ferrara & Taiti, 1986 Somalodilloides pseudopilosus n. sp.
Acknowledgments
SIEGFRIED HUBER (Uhldingen-Mühlhausen/Germany) has donated the isopod material used in this article to the SMNS; Dr. S. Taiti (Florence/Italy) gave advice for the identification of the treated species, he and Dr. S. SFENTHOURAKIS (Nicosia/Cyprus) helped to improve the original manuscript, and Dr. K. Worr- SCHWENNINGER (SMNS) operated the scanning electron micro- scope and the Leica Macroscope. To all of them I wish to express my sincere thanks. Collection permission was granted to Prof. P. WEYGOLDT (no. 07/2000, date 23.1X.2000).
32 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
2 Methods
Photographs were taken through a Leica Z16 APO Macro- scope, processed with Leica Application Suite Version 3.1.8 to obtain combined photographs with extended depth of field; Fig. 1 is a SEM-photograph from air-dried material. The mounted specimen was coated with a 20nm Au/Pd layer and examined with a Zeiss EVO LS15 scanning electron microscope.
Abbreviation
SMNS Staatliches Museum ftir Naturkunde Stuttgart
(+ number of isopod collection), Germany
3 New terrestrial isopod material from Oman
3.1 Chaetophiloscia sp.
Bibliography ScHMALFUuss 1990: 170, figs. 1-17; SCHMALFUss 1991: 2, figs. 4—S.
New material
14,2 29, northern Oman, near Al Hamra, W of Nizwa, leg. S. Huser, IX.2000 (SMNS 11485).
Remarks
Up to now the species of this genus were known from the Mediterranean region. The new record extends the distribution area considerably to the southeast. However, it cannot be excluded that the new record from Oman is due to anthropogenous introduction. The specimens show similarities to C. cellaria, but this taxon seems to consist of a number of separate species. The male is somewhat damaged, so no exact illustrations of the diagnostic char- acters (pleopods) can be given.
Neue Serie 9 3.2 Periscyphis omanensis Taiti & Ferrara, 1991
Bibliography Tarti & FERRARA 1991: 219, figs. 5a-51; Tarri et al. 2000: 147, 152.
New material
1 2, SW-Oman, Dhofar, Tawi Atayr, Wadi Dharbat, +30 km SE of Salalah, leg. S. Huser, X.2000 (SMNS 11483). — 50, 499, SW-Oman, Dhofar, no exact locality, leg. S. Huser, X.2000 (SMNS 11484).—1 &, northern Oman, near Al Hamra, W of Nizwa, leg. S. Huser, IX.2000 (SMNS 11485). —3 SG, north- ern Oman, Al Hamra, W of Nizwa, 23°04'45"N, 57°21'10"E, leg. S. Huser, IX.2000 (SMNS 11485).
Distribution
The species ıs known only from Oman, where it has been found in the northern and in the southwestern parts.
3.3 Xeroniscus erythraeus (Ferrara, 1972) (Fig. 1)
Bibliography
FERRARA 1972: 211, figs. 2-13 (Periscyphis e.); FERRARA & Taırı 1986: 96, figs. 7a-f, 17 (Periscyphis e.); FERRARA & TAITI 1990: 93, figs. 1, 4A—4C.
New material
233, 19, Northern Oman, near Al Hamra, W of Nizwa, leg. S. Huser, IX.2000 (SMNS 11485).
Distribution
The species X. erythraeus was known up to now from northern Ethiopia and from southwestern Saudi Arabia. First record for Oman.
Fig. 1. Xeroniscus erythraeus, 8, 7x3 mm (SMNS 11485), pereopod 7, frontal view, SEM-photograph. — Scale: 0.1 mm.
SCHMALFUSS, NEW SPECIES AND NEW RECORDS OF TERRESTRIAL ISOPODS FROM OMAN 33
3.4 Xeroniscus siegfriedhuberi n. sp. (Figs. 2-12)
Material examined
Holotype: 3, -16x8mm, SW-Oman, Dhofar, Tawi Atayr, Wadi Hinna, +40km NE of Salalah, leg. S. Huser, X.2000 (SMNS T602).
Paratypes: 344, 599, same data as holotype (SMNS 11482),
Derivatio nominis
The species is dedicated to SIEGFRIED HUBER (Uhldingen- Mühlhausen/Germany) who collected a great number of highly interesting isopod samples in many parts of the world and donated them to the SMNS.
Description
Maximum dimensions: 16 x 8mm.
Coloration: Tergal parts yellowish, head with dark grey central part, pereon-tergite 1 with dark anterior and posterior stripe and a medial dark bridge, the remaining pereon-tergites with dark posterior stripes and two rows of dark medial speckles, pleon completely dark except epi- mera, so the animal gives a zebra-like impression.
Cuticular structures: Tergites smooth.
Eyes with around 22 ommatidia arranged in 4 rows (ammonite type). Cephalon with profrons slightly convex, pronounced lateral lobes triangular, interocular line visi- ble except in the middle (Fig. 8). Pereonite 1 with strongly concave posterior margin, frontal corners bent upwards (Figs. 8-9). Posterior margin of pereonite 7 sinuous. Tel- son slightly wider than long, but laterally deeply excavated, thus with a long narrow distal part, apex pointed (Fig. 10). Antennae with first flagellar article twice as long as second.
Male: Pereopods 1-3 with brushes of terminally forked spines on merus and carpus (Fig. 11). Pereopod 7 long and slender, merus proximally with process, ischium ventrally concave, frontally with ridge delimiting a conspicuous depression (Fig. 12). Pleopods 1-5 see Figs. 2-7.
Distribution
Southwestern Oman, only known from the type local- ity in Dhofar.
Differential diagnosis
The ascription of this new species to the genus Xeronis- cus is based on the morphology of the pleopod-exopo- dites. They have the same structure as in other species of Xeroniscus, with respiratory organs in all five exopo- dites (see Figs. 2-7). The body structure is somehow dif- ferent from other species of the genus, with pronounced but interrupted interocular line and deeply rounded con- cavities in the hind margin of the first pereon-tergite (see Figs. 8, 9; compare FERRARA & Taıtı 1990 for figures of these characters in other species of Xeroniscus).
3.5 Xeroniscus troglophilus (Taitı, Ferrara & Davolos, 2000)
Bibliography Tarrı et al. 2000: 154, figs. 4a—k, 8.
New material
1 2, northern Oman, Al Hamra, Al Hota Cave, 23°06'14"N, 57°21'58"E, leg. S. HuBER, IX.2000 (SMNS 11486).
Distribution
The species ıs known only from the type locality ın Oman (Al Hota/Al Fallah Cave system, detailed descrip- tion in HANNA & AL-BELUsHI 1996).
3.6 Somalodilloides pseudopilosus n. sp. (Figs. 13-19)
Material examined
Holotype: d,~5.0 2.3 mm, northern Oman, Al Hamra, W of Nizwa, leg. S. HuBer, [X.2000 (SMNS T603).
Derivatio nominis
The species name indicates the great similarity with S. pilosus Taiti & Ferrara, 2004 (see TAıtı & FERRARA 2004: 298, figs. 58a—h, 59a—h, pl. 25).
Description
Dimensions: ~5.0 x 2.3 mm.
Coloration: White without pigmentation.
Cuticular structures: Tergites thickly covered with apı- cally blunt scale-setae (Fig. 13), which are on posterior margins of tergites not fan-shaped as in S. pilosus; body outline fringed with long pointed setae as in S. pilosus.
Eyes with around 10 ommatidia. Cephalon with thin margin separating frontal shield and vertex. Pereon- epimeron 1 with lateral thickening, dorsally not concave, separated from sulcus arcuatus by a ridge, sulcus arcu- atus extending along epimeron about three quarters of its length, anteriorly not as wide as in S. pilosus (Figs. 13, 15). Posterior margin of pereon-tergite 1 straight. Outer and inner lobe of schisma rounded, the inner one slightly protruding backwards as in S. pilosus. Telson with short distal part and truncate apex as in S. pilosus (Fig. 16). Antenna short and stout, flagellum as long as distal article of peduncle, distal flagellar article three times as long as proximal one. Pereopods with long pointed setae without denticulate apex (Fig. 14). Uropod with minute exopodite, inserted dorsally near posterior margin of protopodite.
Male: Pereopods without sexual modifications. Pleo- pod-exopodite 1 about twice as wide as long with rounded medial part, endopodite I with distal part straight (Fig. 17). Pleopod-exopodite 2 with very long medial part, which is only slightly surpassed by endopodite 2 (Fig. 18).
34 STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
Figs. 2-7. Xeroniscus sieg friedhuberi n. sp., holotype d, 16 x 8mm (SMNS T602). — 2. Pleopod-exopodite 1. 3. Pleopod-endopodites 1, apical parts. 4. Pleopod 2. 5. Pleopod-exopodite 3. 6. Pleopod-exopodite 4. 7. Pleopod-exopodite 5.
SCHMALFUSS, NEW SPECIES AND NEW RECORDS OF TERRESTRIAL ISOPODS FROM OMAN
Figs. 8-10. Xeroniscus siegfriedhuberi n. sp., holotype @, 16 x 8 mm (SMNS T602). — 8. Head and pereon-segment | in frontal view. 9. Head and pereon-segment | in dorsal view. 10. Terminal part of pleon, dorsal view. — Scales: 2mm (8-9), 1 mm (10).
36
STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
Figs. 11-12. Xeroniscus siegfriedhuberi n. sp., holotype 3, 16 x 8mm (SMNS T602). — 11. Pereopod 1, frontal view 12. Pere- opod 7, frontal view. — Scales: 1 mm.
SCHMALFUSS, NEW SPECIES AND NEW RECORDS OF TERRESTRIAL ISOPODS FROM OMAN 37
Figs. 13-14. Somalodilloides pseudopilosus n.sp., holotype 3, ~5.0 x 2.3mm (SMNS T603). — 13. Lateral view of whole animal. 14. Pereopod 7. — Scales: 1 mm (13), 0.2 mm (14).
38 STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
Figs. 15-19. Somalodilloides pseudopilosus n. sp., holotype 4, ~5.0 x 2.3 mm (SMNS T603). — 15. Lateral view of pereon-segment 1. 16. Dorso-caudal view of telson and uropods in situ. 17. Pleopod 1 in ventral view. 18. Apex of pleopod-endopodite 1 enlarged. 19. Pleopod 2 in ventral view.
SCHMALFUSS, NEW SPECIES AND NEW RECORDS OF TERRESTRIAL ISOPODS FROM OMAN 39
Distribution
The new species is known only from the type locality in northern Oman.
Differential diagnosis
Somalodilloides pseudopilosus n. sp. is very similar to S. pilosus from Socotra Island (see Tait & FERRARA 2004: 298, figs. 58a—h, 69). There are, however, some differences which seem to justify the erection of a new species: Sul- cus arcuatus shorter, extending for only two thirds of the length of the epimeron, anteriorly not unusually widened; setae on pereopods with pointed apex, not denticulate as in S. pilosus, male pleopod-exopodite I with rounded medial corner, not angulate as in S. pilosus, width-length relation of pleopod-exopodite 2 is 1.0: 1.6 (in S. pilosus 1.0: 1.1).
4 References FERRARA, F. (1972): The genus Periscyphis Gerstaecker (Crusta-
cea Oniscoidea Eubelidae) in Ethiopia. — Monitore zoologico italiano, Nuova Serie, Supplemento 4: 207-241.
Author’s address:
FERRARA, F. & Taıtı, S. (1986): The terrestrial isopods (Onisci- dea) of the Arabian Peninsula. — Fauna of Saudi Arabia 7: 93-121.
FERRARA, F. & Taıtı, S. (1988): Terrestrial Isopods from Oman (Crustacea). — Journal of Oman Studies, Special Report 3: 391-396.
FERRARA, F. & Tati, S. (1990): A new genus of Eubelidae (Crus- tacea Isopoda Oniscidea) from the Horn of Africa and the Arabian Peninsula. — Tropical Zoology 3: 89-105.
Hanna, S. & AL-Be usu, M. (1996): Introduction to the caves of Oman, 128 pp. (Sultan Qaboos University, College of Science).
ScHMALFuss, H. (1990): Die Landisopoden (Oniscidea) Griech- enlands. 11. Beitrag: Gattung Chaetophiloscia (Philoscii- dae). — Revue suisse de Zoologie 97: 169-193.
ScHMALFuss, H. (1991): The terrestrial isopod genus Chaet- ophiloscia (Oniscidea: Philosciidae) in western Asia. — Stuttgarter Beitrage zur Naturkunde, Serie A, 463: 9 pp.
TAITI, S. & FERRARA, F. (1991): New species and records of ter- restrial isopods (Crustacea) from the Arabian Peninsula. — Fauna of Saudi Arabia 12: 209-224.
TAITI, S. & FERRARA, F. (2004): The terrestrial Isopoda (Crusta- cea: Oniscidea) of the Socotra Archipelago. — Fauna of Ara- bia 20: 211-3235.
TAıTı, S., FERRARA, F. & DavoLos, D. (2000): The terrestrial Iso- poda (Crustacea: Oniscidea) of Oman. — Fauna of Arabia 18: 145-163.
Dr. HELMUT SCHMALFUSS, Staatliches Museum für Naturkunde, Rosenstein 1, 70191 Stuttgart, Germany;
e-mail: schmalfuss.ehrenamt@smns-bw.de
Manuscript received: 15.1V.2015, accepted: 11.V1.2015.
BHL in An
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Stuttgarter Beiträge zur Naturkunde A, Neue Serie 9: 41—62; Stuttgart, 30.1V.2016. DOI: 10.18476/sbna.v9.a4
Two new genera of Caeninae (Ephemeroptera: Caenidae), with a cladistic analysis of the tribe Clypeocaenini
PETER MALZACHER & ARNOLD H. STANICZEK
Abstract
The new genera Mandelara n. gen. (type species M. immutata n. sp.) from South Africa and Papuaenis n. gen. (type species P. balkei n.sp.) from Papua New Guinea are described. Both genera are included in the tribe Cly- peocaenini that is defined by a bulged frons and more or less protruding clypeus. A cladistic analysis recovers the Clypeocaeninae as monophyletic. Caenini + Tasmanocoenini are revealed as sister group to Clypeocanini, together constituting the subfamily Caeninae. A larval key in provided to the genera of Clypeocaeninı.
Key words: Clypeocaenini, new genera, Mandelara, Papuaenis, phylogeny, South Africa, Papua New Guinea.
Zusammenfassung
Zwei neue Gattungen der Familie Caenidae werden beschrieben: Mandelara n. gen. (Typus-Art M. immutata n.sp.) von Südafrika und Papuaenis n. gen. (Typus-Art P. balkei n. sp.) von Papua Neuguinea. Beide Gattungen werden dem Tribus Clypeocaenini zugeordnet, dessen Gattungen sich durch ausgebeulte Frons und mehr oder weniger vortretenden Clypeus auszeichnen. Durch eine kladistische Analyse wird die Monophylie des Tribus abge- sichert. Caenini + Tasmanocoenini bilden die Schwestergruppe der Clypeocaenini, alle zusammen die Unterfami- lie Caeninae. Es wird ein Larvenschlüssel für die Gattungen der Clypeocaenini erstellt.
41
Contents
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1 Introduction
MALZACHER (2013) established the tribe Clypeocaen- ini within the subfamily Caeninae. The Clypeocaenini so far comprise the genera Kalimaenis (Oriental region, two species), Amercaenis (Nearctic region, two species), Cal- listellina (Madagascan region, one species), Trichocaenis (Madagascan region, one species), Provonshara (Ethi- opian region, one species), Clypeocaenis (Ethiopian and Oriental regions, seven species), and Barnardara (Ethi- opian region, one species) (MALZACHER 2013, 2014). All these genera share two synapomorphic characters that define the Clypeocaenini as monophyletic taxon, namely (1) the reduction of filaments on abdominal gill HI to 8 filaments at most, and (2) head bulged in lateral view, clypeus more or less protruding (MALZACHER 2013).
In this contribution, two new monospecific genera, Mandelara n. gen. from South Africa and Papuaenis n. gen. from Papua-New Guinea, are described and placed within Clypeocaenini.
MALZACHER (2013) presented a first phylogeny of Clypeocaenini and improved it with the description of Provonshara (MALZACHER 2014). Both phylogenies were based on traditional Hennigian methodology. In this con- tribution, for the first time a cladistic analysis of Clypeo- caenini is performed to check the monophyly of this taxon and to unravel its intergeneric relationships.
Acknowledgements
We are indebted to MIcHEL Sartori and JEAN-LUC GATTOLLIAT (Musée cantonal de Zoologie, Lausanne) for providing mate- rial to this study, to SUSANNE LEIDENROTH and MILAN PALLMANN (Staatliches Museum ftir Naturkunde, Stuttgart) for mak- ing SEMs and microphotographs, and to MicHEL SARTORI who kindly read the manuscript and provided valuable suggestions.
2 Material and methods The investigated material is preserved in 75% ethanol.
Mandelara specimens were collected by a joint collection of the Musée Cantonal de Zoologie, Lausanne and Albany Museum,
42 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
Grahamstown in the Kruger National Park, South Africa, and that of Papuaenis by the BALKE Expedition 2006 (Zoologische Staatssammlung München) in Papua New Guinea. The holo- types and paratypes are stored in the Musée cantonal de Zoolo- gie, Lausanne.
Specimens used for SEM were dehydrated through a step- wise immersion in ethanol and then dried by critical point dry- ing. The mounted material was coated with a 20nm Au layer, examined and photographed with a Zeiss EVOLS15 scan- ning electron microscope. Macrophotographs were taken with a Leica ZI6APO Macroscope, processed with Leica Applica- tion Suite™ Version 3.1.0 to obtain combined photographs with extended depth of field. All digital photographs were subse- quently enhanced by using PhotoFiltre 7.2.1.
For the cladistic analysis, 14 species of Caenidae were selected, Neoephemera youngi (Neoephemeridae) was chosen
Neue Serie 9
as outgroup (details see under section 5). Seven imaginal and 17 larval characters (see section 5) were chosen and treated as unordered character (except of character 15, which was treated as ordered). The data matrix (Tab. 1) was compiled in Nexus Data Editor (NDE) v.0.5.0 in ss file format converted into NEXUS file format (Mappison et al. 1997), and analysed using TNT v.1.1 (Tree analysis using Traditional Search) (GOLOBOFF et al. 2008). For the Traditional Search, 99999 randomly seeded Wagner trees with 10000 replications were computed with 100 trees to save per replication. All trees found were kept. A strict consensus tree was calculated using all trees. The matrix was resampled with Jacknife 36% removal probability, num- ber of replicates 10000, output results as frequency differences (GC). Consensus tree and Jacknife yielded identical trees. Syn- apomorphies common to all calculated trees were delineated.
Fig. 1. Mandelara immutata n.sp., larva (a—e), Papuaenis balkei n. sp., larva (f-n). — a, f. Sternum IX. b, g. Marginal setation of segment VII. c, h. Marginal setation of segment V. d, I. Hind claw. e, m. Outline of head, lateral view. i. Coxal process of mid leg. j. Coxal process of hind leg. k. Fore claw. n. Lateral outline of pronotum and anterior part of mesonotum.
MALZACHER & STANICZEK, NEW GENERA OF CAENINAE, WITH CLADISTIC ANALYSIS OF THE CLYPEOCAENINI 43
3 Systematic account
3.1 Genus Mandelara n. gen.
Type species: Mandelara immutata n. sp. by present desig- nation.
Etymology
The genus is dedicated to NELSON MANDELA (1918-2013), the late president of the Republic of South Africa.
Differential Diagnosis
Mandelara n. gen. can be characterised and distin- guished from all other genera of Caenidae by the follow- ing combination of characters:
Larva. Great parts of cuticle covered with shield- or funnel-shaped microtrichia (Figs. 6, 7). — Thorax broad- ened (Fig. 2). — Head without ocellar tubercles. — Outline of head with bulges in lateral view (Fig. le). — Clypeus scarcely protruding anteriorly, without setation (Fig. 3). — Maxillar and labial palps three-segmented. — Fore tibia and fore tarsus without filtering setae. — Mid and hind tibia ventrally without long bristles. — Operculate gill ven- trally with an irregular row of simple, scale-shaped micro- trichia, basal half of row also with clusters of spines and numerous single spines (Fig. 5). — Row not reaching hind margin of gill (Fig. 4). — Nearly all filaments of gill III l- or 2-branched, only 1(-3) filaments with 3 or more branches. — Lateral spines of abdomen not bent dorsally. — Posterior part of sternum IX rounded, not shortened, hind margin medially with a small semicircular indentation (Fig. la). — Abdominal terga VII-IX without mediolongi- tudinal ridge. — Hind margin of sternum IX without a bi- pointed process with concave margin between the points.
On the differences to other genera of the Clypeocaen- ini see the larval key below (section 4).
Mandelara immutata n. sp. (Figs. la—e, 2-8)
Holotype, @ larva (microslide): South Africa, Riv. Crocodile/Riv. Incomati, KNP near Malelane, 31°37'04" E, 25°23'57-S) 17.9.2003:
Paratype: 1 larva, same dates as holotype.
Etymology
The species epithet is chosen because of the lack of unam- biguous autapomorphies.
Larva
Measurements and colouration
Subadult female larva, body length 3.3 mm, length of cerciı 2.5mm.
Colouration of cuticle: Intense yellowish-brown, legs and abdomen lighter.
Epidermal pigmentation: Head in front of lateral ocelli dark brownish-grey, with a blackish line between the bases of antenna. Abdominal terga I, HW and VII-IX with broad transverse brownish bands; operculate gills brown- ish, most intense in anterolateral part.
Morphology
Cuticle: Large parts of dorsal and ventral side cov- ered with laminar, shield- or funnel-shaped microtrichia (Figs. 6, 7) (often hardly to be seen in light microscope). Cuticle of abdomen and operculate gill denticulated. Den- ticles of different length and shape and mostly covered by the laminar microtrichia (Fig. 6).
Head: Outline of head bulged ın lateral view (Fig. le). Clypeus scarcely protruding anteriorly; genae moderately bulging out (Fig.3). Mandibles dorso-laterally with bris- tles of different length. Maxillary palp relatively short and broad; segment 3 of labial palp as long as segment 2 (Fig. 3b).
Thorax: Sides of pronotum straight and parallel (Fig. 3a). Legs relatively short and stocky. Transverse row on fore femur consisting of 8—12 spatulate bristles of moderate length. Coxal processes forming inconspicuous ridges. Mid and hind legs with simple bristles of different length more or less blunt, on femora and tibiae nearly all bristles marginally. Fore tarsus with an inner row of about 10-12 strong simple bristles. Mid tarsus with inner row of 10-12 simple bristles and short outer row of 0-4 simple bristles, the apical one unipinnate. Hind tarsus with inner row of 10-12 simple bristles, the apical ones long and slen- der, and an outer row of about 10 small and simple bristles, the apical 4 unipinnate. Claws of all legs slightly bowed, with about 12 denticles (Fig. 1d).
Abdomen: Posterolateral processes short (Fig. 2). Lateral margins with short spatulate or blunt bristles (Fig. la—c). Posteromedian process of tergum II long, tri- angular and pointed, slightly bowed in lateral view. Hind margin of tergum VII with spatulate bristles (Fig. 8a), VIII-X with strong denticles apically more or less rounded (Fig. 8b, c), tergum VIII with a few additional bristles, clearly shorter than those from tergum VII (Fig. 8b). Basal half of tergum VII densely covered with long bipinnate bristles (Fig. 7). Hind margin of sternum IX with a semi- circular indentation, dorsal shagreen not discernible (Fig. la). Gill I nearly half as long as operculate gill. Lat- eral and hind margins of operculate gill with short spat- ulate and pinnate bristles (Figs. 4, 6), inner margin with thin and pointed ones. Medial ridge slightly keeled, with 3—4 short and strong bristles at the base. On ventral side of operculate gill an irregular row of microtrichia running in a greater distance from lateral and hind margin, ending far from the latter (Fig. 4). The row consists of simple scales of different size and shape, in the basal part of the row also with clusters and a broad band of single spines (Fig. 5a, b). Gills III-V) with only 1 filament with 3 branches; all the
44 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
Neue Serie 9
Fig. 2. Mandelara immutata n. sp., larva, light microscope (a), young larva, SEM (b).
other filaments (about 30) simple or two-branched. Basal segments of cerci with apically rounded bristles.
Imaginal stages unknown.
3.2 Genus Papuaenis n. gen. Type species: Papuaenis balkei n. sp. by present designation.
Etymology
The genus name is a combination of Papua (New Guinea) where the genus was found, andCaenis.
Differential Diagnosis
Papuaenis n. gen. can be characterised and distin- guished from all other genera of Caenidae by the follow- ing combination of characters:
Larva. Large parts of cuticle densely covered with shield-shaped microtrichia (Figs. 10, 15, 20). — With broad
spatulate and frayed, sometimes tuft-shaped bristles of dif- ferent length, arranged marginally on femora, tibiae, oper- culate gills etc. (Fig. 21). — Different types of frayed and comb-shaped bristles particularly on maxilla, labrum and fore tibia (Fig. 16). — Thorax broadened (Fig. 9). — Head without ocellar tubercles. — Outline of head with bulges in lateral view (Fig. Im). — Clypeus slightly protruding anteri- orly, without setation (Fig. 10). — Maxillary and labial palps three-segmented. — Fore tibia and fore tarsus without filter- ing setae. — Mid and hind tibia ventrally without numerous long bristles. — Gill I only about 0.2 the length of operculate gill. — Operculate gill ventrally with a row of simple scale- shaped microtrichia of different shape and size (Fig. 19). — Row not reaching hind margin of gill (Fig. 17). — Nearly all filaments of gill III 1- or 2-branched, only 1(-3) filaments with 3 or more branches. — Lateral spines of abdomen not bent dorsally. — Abdominal terga VII-IX with mediolongi- tudinal ridge (Fig. 9). — Posterior part of sternum IX short- ened and cut (Fig. If), without a bi-pointed process with concave margin between the points.
MALZACHER & STANICZEK, NEW GENERA OF CAENINAE, WITH CLADISTIC ANALYSIS OF THE CLYPEOCAENINI
Fig. 3. Mandelara immutata n. sp., larva, head and pronotum (a), mouthparts (b).
45
STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
Figs. 4-5. Mandelara immutata n. sp., larva, operculate gill. — 4. Ventral view. 5. Microtrichia from ventral side, sec- tor from median part of the band (a) and near posterolateral corner (b).
MALZACHER & STANICZEK, NEW GENERA OF CAENINAE, WITH CLADISTIC ANALYSIS OF THE CLYPEOCAENINI
Figs. 6-7. Mandelara immutata n. sp., larva. — 6. Operculate gill, microtrichia on dorsal side. 7. Sector from tergum VII.
47
Neue Serie 9
STUTTGARTER BEITRAGE ZUR NATURKUNDE A
48
Fig. 8. Mandelara immutata n. sp., larva, sector from hind margin of tergum VII (a), VIII (b) and IX (c).
MALZACHER & STANICZEK, NEW GENERA OF CAENINAE, WITH CLADISTIC ANALYSIS OF THE CLYPEOCAENINI 49
Egg. Epithemata very large, with single threads with tassel-shaped terminal knobs running along the meridians (Figss22 223).
On the differences to other genera of the Clypeocaen- ini see the larval key below (section 4).
Papuaenis balkei n. sp. (Figs. 1f—n, 9-24)
Holotype, 6 larva (microslide): PNG 102, Papua New Guinea, Marobe, Wagau, Herzog Mts., 146°48'068" E, 06°51'067" S, 19.X1.2006, BALKE & KINIBEL leg.
Paratypes: PNG 87, Papua New Guinea, Eastern Highlands, Marawaka, Ande, 145°49'807"E, 07°01'697"S, 09.X1.2006, 192 larva, BALKE & KuiniBeL leg. — PNG 90, Papua New Guinea, Gulf, Marawaka, Mala, 145°44'467" E, 07°05'664" S, 11.X1.2006, 2 99 larvae, BALKE & KINIBEL leg. — PNG 119, Papua New Guinea, Madang, Keki, Adalbert Mts., 145°24'437" E, 04°43'058" S, 29.X1.2006, 19 larva, BINATANG Boys.
Etymology
The species is dedicated to the collector and leader of the Papua New Guinea expedition 2006, MICHAEL BALKE (Zoologi- sche Staatssammlung München).
Figs. 9-11. Papuaenis balkei n. sp., larva. — 9. Habitus. 10. Head and pronotum. 11. Maxilla.
50 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
Larva
Measurements and colouration
Subadult male larva, body length 3.5mm, length of cerci 1.5 mm; subadult female larva, body length 4.2 mm, length of cerci 2.2 mm.
Colouration of cuticle: Beige. Head, pro- and mesono- tum, operculate gills, and femora with a brownish shad- ing, caused by the brown microtrichia.
Epidermal pigmentation: No pigments visible.
Morphology
Cuticle: Large parts of dorsal side densely covered with small, compact, shield-shaped microtrichia (Fig. 20). A conspicuous pattern of pale marks, blotches, and lines results from the lack of microtrichia on these places (Figs. 9, 10).
Head: Outline of head bulged in lateral view (Fig. Im), clypeus slightly protruding anteriorly; genae strongly bulging out (Fig. 10). Mandibles with only one or two long bristles and a ventrobasal field densely covered with broad multi-branched bristles (as in Fig. 21a). Galealacinia apico-medially broadly rounded (Fig. 11). Segment 3 of maxillary palp ventrally with numerous very long, thin and medially shortly combed bristles; peeking out beneath them some broad comb-shaped bristles with very long and thin tips (Fig. 12). Segment 3 of labial palp about 0.8 times as long as segment 2 (Fig. 13). Postmentum with similar bristles as on maxilla (Fig. 21a).
Thorax: Sides of pronotum straight and parallel, some- times fore corner laterally a little protruding and side slightly S-shaped; sides of mesonotum with broad rounded antero-lateral projections (Figs. In, 9). Femora broad and flattened (Fig. 14). Fore femur with a very dense transverse row of 15-18 bristles which are broadly spatulate, bifur- cated and pinnate (Figs. 14a, 15). Coxal processes flatly semi-elliptical and slightly bulging out (Fig. 11, j). Mid and hind legs marginally with similar bristles like that from the transverse row on fore femur (Fig. 14b). Tibiae with comb-shaped, unipinnate bristles (Fig. 16b), apically addi- tionally with strongly bipinnate ones (Fig. 16a). There are a lot of different shapes from short and very broad bristles to those of moderate length and also transitional stages to tuft-shaped forms (fore tibia) (Fig. 21). Fore tarsus with an inner row of about 8 relatively long simple bristles. Mid tarsus with inner row of about 7 strong simple bris- tles. Hind tarsus with inner row of about 10 strong simple or slightly unipinnate bristles and an outer row of about 5 small and unipinnate bristles. Claws short and basally broad, with sides clearly converging anteriorly; apically bent, without denticles (Fig. Ik, 1).
Abdomen: Posterolateral processes of moderate length (Fig. 1f-h). Posteromedian process of tergum II broadly triangular with rounded tip. Terga VII-IX with medio-
Neue Serie 9
longitudinal ridge (Fig.9). Hind margin of tergum VII with short spatulate and pinnate bristles, VIII—X with long denticles apically more or less rounded. Tergum VII cov- ered with spatulate bifid bristles that are slightly pinnate. Posterior part of sternum IX shortened and cut, sometimes medially slightly indented (Fig. If), dorsal shagreen not discernible. Gill I very short, about 0.2 the length of oper- culate gill. Lateral and hind margins of operculate gill with broadly spatulate and pinnate bristles (Fig. 18), inner mar- gin with thin, pointed and finely frayed ones. Y-shaped ridge keeled, the inner branch a little stronger, with 3—4 short and strong bristles at the base. Row of microtrichia on ventral side of operculate gill running in regular dis- tance from the lateral and hind margins, not reaching the latter (Fig. 17). The row consists of short and rounded sim- ple microtrichia (Fig. 19). Gills III-V) with only 1 fila- ment with 3 branches; all the other filaments simple or two-branched.
Eggs
[The eggs were taken from a last instar female larva. In this developmental stage, parts of the egg surface, par- ticularly the epithemata, are still covered by a thin protect- ing membrane. ]
Eggs short oval or barbell-shaped. Chorion with irreg- ularly distributed, wide pores (Fig. 24). Two large epithe- mata; except for a narrow equatorial area covering the whole egg surface (Figs.22, 23), not corresponding to types of epithemata hitherto described for Caenidae. A large number of single threads, each with a terminal tas- sel-shaped knob, running from the polar region along the meridians towards the equator (single-thread-type). After dissolution of the protecting membrane threads float- ing in the surrounding water (Fig. 23b). One (or two?) micropyle(s) with a short or very short visible channel and a small, irregular sperm-guide (Fig. 24).
Imaginal stages unknown.
4 Larval key to the genera of Clypeocaenini
1 Fore tibia and fore tarsus with long filtering setae.............. 2
— Fore tibia and fore tarsus without filtering setae................. 5
2 Filtering setae reaching to about half the length of tibia, densely arranged medially on fore tibia and fore tarsus. ......
ee wn Bb he Wath eet N. ER aene Amercaenis — Filtering setae nearly as long as tibia, not densely arranged and also present laterally on tibia and tarsus....................... 3
3 Maxillary palp three-segmented. Body cuticle with strong long spines. Mid and hind tibia ventrally with numerous long bristles (MALZACHER 2014: fig. Ic). ........... Provonshara
— Maxillary palp two-segmented. Body cuticle without strong long spines. Mid and hind tibia ventrally without long DEISTIES. A 2, saan Ria Paths ot ae ee eas aaa aa 4
MALZACHER & STANICZEK, NEW GENERA OF CAENINAE, WITH CLADISTIC ANALYSIS OF THE CLYPEOCAENINI
a FP. ee
(da
„ala ee a aac,
i — Tr Mille. ree LTT Tr ny, ‘ a ae Er TI ms 5 ° er, ty a
Figs. 12-13. Papuaenis balkei n.sp., larva. — 12. Comb-shaped bristles from segment 3 of maxillary palp. 13. Labium.
51
52
STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
A
Figs. 14-15. Papuaenis balkei n. sp., larva. — 14. Fore leg (a) and hind leg (b). 15. Sector from transverse row of bristles on fore femur.
MALZACHER & STANICZEK, NEW GENERA OF CAENINAE, WITH CLADISTIC ANALYSIS OF THE CLYPEOCAENINI
Figs. 16-17. Papuaenis balkei n. sp., larva. — 16. Different bristles from fore tibia, apical protrusion (a), inner margin (b). 17. Operculate gill, ventral view.
53
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Figs. 18-19. Papuaenis balkei n. sp., larva, operculate gill. — 18. Bristles from lateral margin. 19. Microtrichia from ventral side, sector from median part (a) and from basal part (b) of the row.
MALZACHER & STANICZEK, NEW GENERA OF CAENINAE, WITH CLADISTIC ANALYSIS OF THE CLYPEOCAENINI
Figs. 20-21. Papuaenis balkei n. sp., larva. — 20. Microtrichia from dorsal cuticle (a), in higher magnification (b). 21. Different bristles, from postmentum (a), fore tibia (b) and fore femur (c).
55
STUTTGARTER BEITRAGE ZUR NATURKUNDE A
Filtering setae forming regular rows (PROVONSHA & McCarrerty 1995: fig. 16). Clypeus anteriorly strongly pro- truding: “with lotissetac) aller me Clypeocaenis Filtering setae irregularly arranged (PROVONSHA & McCarrerty 1995: fig. 5). Clypeus anteriorly without long SCLAG.: Se Leta de A En. uR Lens Tale tester, rue A Barnardara Legs long and slender, femora narrowed (MALZACHER 2013: fig. 8b-d). Maxillary palp elongated and coiled (MALZACHER 2013: fig. 12a). Mesonotum not broadened......... Kalimaenis Legs not elongated, femora broad. Maxillary palp shorter and not coiled. Mesonotum broad, outline of body more or lesseveniyzcunrvedieerar u Maar fry are ee 6 Great parts of cuticle densely covered with shield-shaped TIErOLLIc Ha. IE SE6R2 0) er A en nn en 7 Cuticle without densely arranged shield-shaped microtri- CUNT ant Sa AR ARTE RR | 8 Sternum IX posteriorly shortened, hind margin straight (Fig. If). Claws short, without denticles (Fig. 1k-l)............... RR EEE RE ET ER Paint eg aA ee, me Papuaenis Sternum IX not shortened, hind margin bowed, with median incision (Fig. la). Claws longer, with about 12 denticles (SSH). Wi Banner a OPE leat Je Mandelara
8
Neue Serie 9
Figs. 22-24. Papuaenis balkei n. sp., eggs. — 22. Barbell-shaped egg. 23. More rounded egg (a), floating threads (b). 24. Micropyle.
Body surface provided with many very long hair-like bris- tles (MALZACHER 2009a: fig. 4). Thoracic notae smooth........ Re en ee Trichocaenis Body surface without long bristles. Thoracic notae with ridges or bulges (Sun & McCarrerty 2001: fig. ])................ IR epee > Ude. IERER Abi 5ER Mehrere. Se N er Callistellina
5 Cladistic analysis
For data matrix see Tab. 1. Characters used for cladis-
tic analysis are as follows:
1 2 B
4
(1) Mesonotal ommation: (0) absent; (1) present.
(I, L) Hind wing (buds): (0) present; (1) absent.
(1) Segmentation of forceps: (0) 4-segmented; (1) l-segmented.
(I, L) Styliger (anlagen): (0) fully exposed; (1) partly retracted into abdominal segment IX.
(I) Procoxae: (0) broadly separated; (1) approximated.
MALZACHER & STANICZEK, NEW GENERA OF CAENINAE, WITH CLADISTIC ANALYSIS OF THE CLYPEOCAENINI 57
Tab. 1. Data matrix for 14 species of Caenidae and Neoephemera youngi (Neoephemeridae).
= 5 5 = 5 5
EM ea ail ts |
1 1 1 I 1
6 (I) Metathorax: (0) without transversal ridge; (1) with transversal ridge.
7 (I) Shape of forceps: (0) without ridges or keels; (1) with longitudinal ridge; (2) with single ventral longitu- dinal groove; (3) with dorsal and ventral longitudinal groove.
8 (L) Gill cover medially: (0) not overlapping; (1) over- lapping.
(L) Gill cover: (0) bilobed; (1) without ventral lobe.
10 (L) Ventral side of gill cover: (0) without microtrichia; (1) with field of irregularly arranged microtrichia; (2) with band of irregularly arranged microtrichia; (3) with single row of regularly arranged microtrichia; (4) with transverse rows of regularly arranged micro- trichia.
11 (L) Gill III with multiple branchings: (0) in nearly all filaments; (1) in 15-25 filaments; (2) in about 8 fila- ments; (3) in 1-3 filaments.
12 (L) Abdominal posterolateral projections in abdominal segments: (0) (2)3-9; (1) 4-7; (2) 3-7, (3) 2, 5-9.
13 (L) Femora: (0) broad and stout; (1) thin and elongated; (2) extremely broadened and stout.
14 (L) Profemora dorsally: (0) with transverse row of setae; (1) without transverse row of setae.
15 (L) Protibiae: (0) without filter setae; (1) with irreg- ularly arranged short filter setae; (2) with irregularly arranged long filter setae; (3) with regularly arranged rows of filter setae.
16 (L) Outline of head: (0) evenly bowed; (1) bulged.
17 (L) Clypeus anteriorly: (0) not protruding; (1) slightly protruding; (2) heavily protruding.
18 (L) Maxillary palps: (0) 3-segmented; (1) 2-segmented; (2) 1-segmented.
19 (L) Labial palps: (0) 3-segmented; (1) 2-segmented.
olılılılalzlılo/ılololojolo/ılılılolo|
3)2/olo/olılılılolololı)2lılo,
3)ılolololoJolojoJolo/2]ılolo (3l1ılojololojojojojojo]2]ılolo
Hae EEE 2 EZ ER EI DZ EZ 11
20 (L) Ocelli: (0) without tubercles; (1) with small tuber- cles; (2) with large tubercles.
21 (L) Operculate gill, ventral row of microtrichia: (0) lacking; (1) not reaching hind margin; (2) reaching hind margin.
22 (L) Operculate gill, medial ridge on dorsal side: (0) absent; (1) not keeled; (2) keeled; (3) extremely keeled.
23 (L) Mesonotum: (0) not broadened, laterally with dis- tinct angle to abdomen; (1) broadened, laterally con- fluent with abdomen.
24 (L) Shield-shaped microtrichia on cuticle: (0) absent; (1) present.
6 Phylogenetic discussion
The cladistic analysis confirmed the monophyly of Caenidae, which was unchallenged before. Within Cae- nidae, the Brachycercinae + Madecocercinae were recov- ered as sistergroup to the monophyletic Caeninae. Within Caeninae, the Caenini + Tasmanocoenini and Clypeocae- nini were recovered as well-supported sistergroups.
Within Clypeocaenini, Kalimaenis sibylliana branches off first, followed by Amercaenis ridens. The remaining genera within Clypeocaenini form a weakly supported monophyletic group. Within this group, however there are two well supported monophyla, namely Papuaenis balkei + Mandelara immutata and Provonshara spinifera + (Clypeocaenis oligosetosa + Barnardara demoori). The positions of Callistellina panda and Trichocaenis inex- perta in relation to these other two groups remain unre- solved (see Fig. 25). For apomorphies determined ın the cladistic analysis see Fig. 26.
58 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
Neoephemera youngi
Neue Serie 9
Caenoculis bishopi
AO
Brachycercus harrisella
Madecocercus tauroides
100
50 7 Tasmanocoenis tillyardi
Caenis macrura
Callistellina panda
40 Kalimaenis sibylliana 74 Amercaenis ridens
24 5
Trichocaenis inexperta
Papuaenis balkei ce BR 52
Mandelara immutata Provonshara spinifera
Clypeocaenis oligosetosa 61 Eu yp g
Barnardara demoori
Fig. 25. Cladistic analysis of Clypeocaenini, Strict Consensus, Jacknifing (P=36). — GC Values, 10000 replicates, cut=1 (tree 0).
The cladistic analysis confirms the mental analysis due to Phylogenetic Systematics sensu HENNIG (1966) that results in the very same phylogeny (based on larval char- acters).
Due to mental analysis (Fig. 27), the genera of the tribe Clypeocaenini share two synapomorphies:
(1) Gill II (first of the four respiratory gills) pos- sesses at most eight filaments (in most cases only one or two), each made up of three or more branches, in contrary to 15-25 filaments in the Caenini and Tasmanocoen- ini (character 11). In the outgroup Neoephemeridae there are numerous filaments present, each with 5 and more branches, so that a reduction of filament branches can be regarded as apomomorphic for Clypeocaeninı.
(2) Outline of head in lateral view with bulges, clypeus more or less protruding (character 14, see also MALZACHER 2013: fig. 13c—h). In the above mentioned outgroups and in Ephemerellidae the outline of the head in lateral view is evenly bowed (MALZACHER 2013: fig. 13a, b).
The sister group of Clypeocaenini, represented by the tribes Caenini + Tasmanocoenini shares two synapomorphies:
(3) Row of microtrichia on ventral side of gill II (oper- culate gill) extends to hind margin of gill (character 21; for Caenini see MALZACHER 2009b: figs. 17, 20 and 23; for Tasmanocoenini see SUTER 1984: fig. 6, ALBA-TERCEDOR & SuTER 1990: fig. 19, SUTER 1993: figs. 16, 41 and 68, and Suter 1999: fig. 95). In Clypeocaenini, the row of micro- trichia does not extend to hind margin of operculate gill. The distance from the end of the row to hind margin varies from '/s to '/is the length of gill II (Figs. 4, 17).
(4) Gill II with regular row of complex, very similar, scale-shaped microtrichia, which are semicircular or more or less elongated, consisting of 20-30, rather basally fused filaments or spines (character 10). In Kalimaenis, a simi- lar arrangement is present, but this is regarded as a conver- gent development. The plesiomorphic character states in Clypeocaenini show very different developmental stages from a band of simple spines and clusters of spines up to scales with 5—20 filaments arranged in an irregular row
MALZACHER & STANICZEK, NEW GENERA OF CAENINAE, WITH CLADISTIC ANALYSIS OF THE CLYPEOCAENINI 59
Neoephemera youngi
9,19 Caenoculis bishopi 12,17,18,19 Brachycercus harrisella 12,17 Madecocercus tauroides
6,13
0 Nine apna i tillyardi Caenis macrura 12-Kalimaenis sibylliana Amercaenis ridens
4,5 15,16,21 22
Callistellina panda 9,21-Trichocaenis inexperta 9,104.92 Pee balkei Mandelara immutata
Provonshara spinifera 1 a 14,16,21-Clypeocaenis oligosetosa
Barnardara demoori
Fig. 26. Apomorphies of cladistic analysis. — Synapomorphies common to three trees.
(MALZACHER 2009b: figs. 3-5, 8, 10, 12). The most plesi- omorphic state, simple spines and clusters of spines, can also be found in Brachycercinae and Madecocercinae. In the outgroup Neoephemeridae there are no ventral micro- trichia present at all.
Within Clypeocaenini the genus Kalimaenis branches off from the remaining genera that share a single apomor- phic character (character 23):
(6) Mesonotum broad. Pronotum broadly attached to the mesonotum (Figs. 2, 9). Outline of body more or less evenly curved. (e. g. MALZACHER 2013: fig. 47, MALZACHER 2009a: fig.4; Sun & McCarrerty 2001: fig. 1). In Kali- maenis, the pro- and mesonotum is much narrower. In MALZACHER 2013 and 2014 this was established errone- ously also for Amercaenis, but it turns out that the mes- onotum of the latter is clearly broader.
Kalimaenis shows three autapomorphies:
(8) Legs long and slender (character 13), particularly in the females, with narrow femora more or less parallel- sided (MALZACHER 2013: fig. 8b-d). This shape shows a
certain similarity to legs of Brachycercinae. The fore legs of Kalimaenis are reduced in length just like in the latter subfamily.
Two further characters were not included in the cladis- tic analysis, but are clearly autapomorphies of Kalimaenis:
(9) Maxillary palps strongly elongated and narrowed with a conical segment 1 and S-shaped coiled segments 2 and 3 (MALZACHER 2013: fig. 12a). As far as we know, this shape is unique in the Caenidae or even in the Pan- nota, where a tendency of reduction can be observed in all families. Only in Tricorythus a long and slender maxil- lary palp can be observed, but it shows only two segments (MCCAFFERTY & WANG 2000: fig. 85).
(10) Hind claws with about seven groups of micro-den- ticles. The micro-denticles in each group are fused together to larger denticles, respectively (MALZACHER 2013: fig. 8e, f). In the Caeninae there can be homodont or heterodont hind claws present. The former are equipped with a num- ber of denticles equal in size, whereas the latter have an additional row of more or less fused microdenticles (e. g.
60 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
MALZACHER 2009a: figs. Ip, 2h, 3g), a state that is apomor- phic in some Caenini but cannot be found in the Clype- ocaenini. On the other hand there is at least one Caenis species that shows a long row of fused microdenticles only (MALZACHER, In preparation). Nevertheless groups of fused micro-denticles are unique in Kalimaenis.
In the next step Amercaenis branches off from the Clypeocaenis group s. str. that shares the following apo- morphy:
(5) In gill III nearly all filaments (up to 45) have one or two branches (character 11). Only one or two apical fil- aments show three or more branches. These are situated at the apex of the gill (MALZACHER 2014: fig. Ik). In Kali- maenis and Amercaenis there are 7—8 filaments with up to 5 branches and only about 20 reduced ones.
£ 2 2 = Be 5 a ee es ee S se @ <6 2 BR “GS DB eS oF 2 „un Fs 8.8 £6 5 8 DE ee u Te 48 8 GE © 3 @ € ¢€ a SG = = SS 22] [46] [rs 14) 10 18] [7 13 9 8 21 12 u 20 19) 5 ie 2 EI 11 3
Fig. 27. Phylogenetic tree concluded by mental analysis with indicated apomorphies (see section 6 for detailed discussion on characters).
Neue Serie 9
Amercaenis shows the apomorphic character:
(19) Densely arranged short filtering setae medially on tibia and tarsus of fore leg. This arrangement can only be found in Nearctic species and represents a parallel develop- ment to the African and Oriental genera with filtering setae.
The remaining genera of Clypeocaenini split into four groups: Trichocaenis, Callistellina, Mandelara + Papuae- nis, and Provonshara + (Barnardara + Clypeocaenis). The phylogenetic relationship of these four groups to each other remains unresolved.
All Clypeocaenini except for Trichocaenis show the character:
(7) Gill II with a keeled inner ridge (character 22; see also MALZACHER 2013: figs. 9d, 43, and PROvoNSHA & MCCAFFERTY 1995: fig. 7). A keeled ridge is also present in some taxa of Caeninı. It may be possible that this charac- ter has evolved independently in these groups, or it is even possible that it is present ın their common groundplan, but reduced in Trichocaenis and most Caenini.
Trichocaenis shows the following apomorphic character:
(20) Body surface provided with a lot of very long hair- like bristles. This character is unique within the Clypeo- caenini and otherwise can only be found in a few Caenını. Thus we conclude that this character has evolved indepen- dently in Trichocaenis and a few Caenini.
Callistellina is well-defined by the following two apo- morphies:
(15) Head with microscopic pits each with a hand- shaped microtrichium (Sun & McCarrerty 2001: figs. 9 and 10; MALZACHER 2009a: fig. 10).
(16) Thoracic notae with ridges or bulges (SUN & McCarrerty 2001: fig. 1). A single medio-longitudinal ridge is also visible in Kalimaenis (MALZACHER 2013: fig. 32).
Mandelara and Papuaenis share the following synapo- morphic character:
(21) Great parts of cuticle densely covered with shield- shaped microtrichia often overlapping themselves with their margins (Figs. 6, 20). Shield or funnel-shaped microtrichia seem to have developed independently in several Caenocu- lis (MALZACHER, In preparation) and Caenis species.
Papuaenis shows one apomorphic character:
(22) Sternite IX posteriorly shortened, hind margin straight, not projecting beyond the posterolateral pro- cesses anteriorly. Such a shortening cannot be observed in other Caenidae.
At present, Mandelara cannot be defined by derived characters.
The Clypeocaenis group with the genera Clypeocae- nis, Barnardara and Provonshara shares the following synapomorphy:
MALZACHER & STANICZEK, NEW GENERA OF CAENINAE, WITH CLADISTIC ANALYSIS OF THE CLYPEOCAENINI 61
(11) Filtering setae on tibia and tarsus of fore leg. This is a parallel development to similar structures in Amercae- nis (see character 9), but filtering setae are not arranged as densely as in Amercaenis and can also be found on the lateral part of tibia and tarsus, reaching more or less the length of tibia (SoLDAN 1978: fig. 8, PROVONSHA & MCCArFFERTY 1995: figs. 5, 16). In Amercaenis, on the other hand, filtering setae reach only a little more than half the length of tibia (PRovoNsHA & McCarrerty 1985: fig. 6, and 2006: fig. 5).
Provonshara is the sister to Clypeocaenis + Barnard- ara. The latter are united by a single apomorphy:
(12) Maxillary palp two-segmented. This character can also be observed as parallel development in the Brachycer- cinae and some families of the Pannota (see MCCAFFERTY & Wang 2000 and KLUGE 2004).
Barnardara is not defined by unique apomorphic char- acters, while Clypeocaenis shows two apomorphies:
(13) Filtering setae arranged in rows. Few setae also often on inner margin of femur and in a basket-like arrangement on mandible (SoLDAN 1978: fig. 8, PROVONSHA & McCarrerty 1995: figs. 13, 16).
(14) Clypeus strongly protruding and provided with long setae.
Provonshara has two unique apomorphies:
(17) Body cuticle with strong long spines (MALZACHER 2014: fig. 2c). In all other Caeninae the cuticle is smooth, granulated, or shows short microdenticles (in Kalimaenis strongly granulated or scaly).
(18) Mid and hind tibia with numerous long bristles on ventral side (MALZACHER 2014: fig. Ic). In other genera those bristles are not present. If at all, long bristles occur only on dorsal side or on the margins of tibia.
It is noteworthy that both cladistic and mental analyses end up with identical phylogenetic hypotheses. In sum- mary, it can be concluded that Clypeocaenini form a well- defined monophyletic group within Caeninae.
7 References
ALBA-TERCEDOR, J. & SuTER, P. J. (1990): A new species of Caeni- dae from Australia: Zasmanocoenis arcuata sp. n. (Insecta, Ephemeroptera). — Aquatic Insects 12: 85-94.
GOLOBOFF, P. A., Farris, J.S. & Nixon, K. C. (2008): TNT, a free program for phylogenetic analysis. — Cladistics 24: 774-786.
HEnnIG, W. (1966): Phylogenetic Systematics, 263 pp.; Urbana, Illinois (University of Illinois Press).
Krug, N. Yu. (2004): The phylogenetic system of Ephemer- optera, XIII + 442 pp.; Dortrecht/Boston/London (Kluwer Academic Publishers).
Mappison, D. R., Sworrorb, D. K. L. & Mappison, W. P. (1997): NEXUS: an extensible file format for systematic informa- tion. — Systematic Biology 46: 590-621.
MALZACHER, P. (2009a): New larvae of Caeninae from Madagas- car (Ephemeroptera: Caenidae). — Stuttgarter Beitrage zur Naturkunde A, Neue Serie 2: 177-194.
MALZACHER, P. (2009b): Comparative morphology of gill cover microtrichia in the Caenidae (Insecta: Ephemeroptera). — In: STANICZEK, A. H. (ed.): International Perspectives in Mayfly and Stonefly Research. Proceedings of the 12'" International Conference on Ephemeroptera and the 16" International Symposium on Plecoptera, Stuttgart 2008. — Aquatic Insects 31, Supplement 1: 479-488.
MALZACHER, P. (2013): Caenidae from East Kalimantan, Borneo (Insecta: Ephemeroptera). With a discussion on phylogeny of the new tribe Clypeocaenini, subfamily Caeninae. — Stuttgarter Beiträge zur Naturkunde A, Neue Serie 6: 21-55.
MALZACHER, P. (2014): A new genus of the tribe Clypeocaen- ini (Ephemeroptera: Caenidae: Caeninae). — Stuttgarter Beiträge zur Naturkunde A, Neue Serie 7: 1-7.
MCCAFFERTY, W.P. & Wang, T.-Q. (2000): Phylogenetic sys- tematics of the major lineages of pannote mayflies (Ephemeroptera: Pannota). — Transactions of the American Entomological Society 126: 9-101.
ProvonsHa, A. V. & McCarrerty, W. P. (1985): Amercaenis: new Nearctic genus of Caenidae (Ephemeroptera). — Interna- tional Quarterly of Entomology 1: 1-7.
ProvonsHa, A. V. & McCarrerty, W. P. (1995): New brushlegged caenid mayflies from South Africa (Ephemeroptera: Caeni- dae). — Aquatic Insects 17: 241-251.
PROVONSHA, A. V. & McCarrerty, W. P. (2006): A second species of the North American mayfly genus Amercaenis Provon- sha and McCafferty (Ephemeroptera: Caenidae). — Journal of Insect Science 6(10): 1-6.
SoLDAN, T. (1978): New genera and species of Caenidae (Ephemeroptera) from Iran, India and Australia. — Acta entomologica bohemoslovaca 75: 119-129.
Sun, L. & McCarrerty, W. P. (2001): Callistina panda, a striking new genus and species of Caeninae (Insecta: Ephemero- ptera: Caenidae) from Madagascar. — Bulletin de la Société d’ Histoire naturelle de Toulouse 137: 7-15.
Suter, P. J. (1984): A redescription of the genus Tasmanocoenis Lestage (Ephemeroptera: Caenidae) from Australia. — Trans- action of the Royal Society of South Australia 108: 105-111.
Suter, P.J. (1993): Wundacaenis, a new genus of Caenidae (Insecta: Ephemeroptera). — Invertebrate Taxonomy 7: 787— 803.
Suter, P.J. (1999): Illustrated key to the Australian cae- nid nymphs (Ephemeroptera: Caenidae.). — Co-opera- tive Research Centre of Freshwater Ecology. Identification Guide 23: 36 pp.
62 STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
Authors’ addresses:
Dr. PETER MALZACHER, Friedrich-Ebert-Straße 63, 71638 Ludwigsburg, Germany; e-mail: malzacher.Ib@t-online.de
Dr. ARNOLD H. STANICZEK, Department of Entomology, State Museum of Natural History, Rosenstein 1, 70191 Stuttgart, Germany; e-mail: arnold.staniczek@smns-bw.de
Manuscript received: 30.1.2015, accepted: 3.1X.2015.
Stuttgarter Beiträge zur Naturkunde A, Neue Serie 9: 63-69; Stuttgart, 30.1V.2016. DOI: 10.18476/sbna.v9.a5 63
Two new Caenis species from north-eastern China (Insecta: Ephemeroptera)
PETER MALZACHER
Abstract
The new mayflies Caenis wui n. sp. and C. pekinensis n. sp. from China (Beijing) are described and compared with West and Central Palaearctic species (C. robusta and species ofthe C. beskidensis group).
Keywords: Caenidae, Caenis, Palaearctic, Beijing, ULMER collection.
Zusammenfassung
Zwei neue Eintagsfliegenarten aus China (Peking), Caenis wui n. sp. und C. pekinensis n. sp., werden beschrie- ben und mit west- und zentralpaläarktischen Arten verglichen (C. robusta und Arten der C. beskidensis-Gruppe).
Contents
-Introduchon wert tr. Feel a he ee er Br 2: Material anchinethod sever of di. ren rs nee ESYSIEINALICACKOUNT Mee ce Aes 8 ote rl ee en on oot RE CACHES WHE NGS Dna een. aan 3:2 Gaenis pekinensis NESPw u... A INETOECIICOS Sc A rn
1 Introduction
Within the bounds of an investigation of the Caenidae in the collection GEORG ULMER (1877-1963) deposited in the Museum of the University of Hamburg, I examined also samples with Caenis specimens from the surround- ings of Beijing, collected by C.F. Wu. In two cases vis- ible subimaginal genitalia allow the description of new species. From China, six species of Caenis are so far described (ZHou & ZHENG 2004, Jia et al. 2010), but they can be well distinguished from the herein described spe- cies. Unfortunately no dates are given on the labels, but it seems very likely that the material described below was part of the material which is mentioned by ULMER (1936). ULMER states that mayflies in alcohol were sent to him by Wu “in den letzten Jahren” [during the last years], that the collector donated the majority of the material to him, and that part of it remained unidentified. Most of Wu’s mate- rial is cited without exact calendar dates also in ULMER’S article, but at least in two cases the years 1929 and 1930 are given. Wu has worked in Beijing since 1926.
Acknowledgements
My special thanks go to the colleagues from the Zoologis- ches Museum Hamburg, in particular to Kar ScHÜTTE for leav- ing me the material for investigation. Thanks also to SUSANNE LEIDENROTH for making the SEMs and to ArnoLD H. STANICZEK (both Staatliches Museum für Naturkunde, Stuttgart) for useful comments on the manuscript.
ER ese etre HR RU EAST, Sec LO aE RMT, MORE Me u et ATED De 66 sheers ROM! sa B ee Zev eR eto O BER Ri eRe sah Vea! Desheean te SE ial eee 68
2 Material and methods
The examined material is preserved in 75% ethanol. The types of the herein described species are deposited in the Zoolo- gisches Museum, Hamburg.
Specimens used for SEM were dehydrated through a step- wise immersion in ethanol and then dried by critical point dry- ing. The mounted material was coated with a 20nm Au layer, examined and photographed with a Zeiss EVO LS 15 scanning electron microscope. Digital photographs were enhanced by using Photofiltre 6.5.2 (http://www.photofiltre-studio.com).
3 Systematic account
3.1 Caenis wui n. sp. (Figs. 1, 3-6)
Holoty pe, d larva (on microslide); M22, China, Peking [= Beijing env.], May [exact date missing], leg. C. F. Wu.
Paraty pes: same data as holotype, 19 larva (differential diagnostic characters are the same in male and female larvae).
Etymology
The new species is dedicated to the collector, CHENFU F. Wu [= Hu Jincru] (1896-1972).
Male imago
The holotype is a male last instar larva. The follow- ing subimaginal features are therefore visible and can be
64 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
Neue Serie 9
Fig. 1. Caenis wui n. sp., larva (a—e) [genitalia of 4 subimago (b) and lateral processes of subimago (b-e) visible], male subimago (f-g). — a. Operculate gill, general view. b. Sternum IX. ce. Abdominal segment VII, lateral part. d. Abdominal segment V, lateral part. e. Abdominal segment III, lateral part. f. Prosternal triangle. g. Forcipes.
described: genitalia, arrangement of prosternal ridges, shape of antenna, and posterolateral processes on abdo- men:
Base of antennal flagellum not dilated; pedicel more than three times as long as scape. Prosternal ridges form- ing a triangle with straight or slightly concave lateral sides and broadly rounded tip (Fig. If). Abdominal segments V-VIII with long or very long lateral processes (Fig. Ic, d), segments III, IV and IX with shorter ones (Fig. 1b, e). Sternite IX and subimaginal genitalia as in Fig. Ib. Penis anvil-shaped. Forcipes moderately short with converging sides and a sclerotized apical knob or rounded tip (Fig. 1g).
Egg [extracted from female last instar larva]
Egg oval, relatively short and broad (Fig. 3). Cho- rion granulated, with two coiled rope-like epithemata of robusta subtype. 15-30 threads emerge from the poles (Fig.4). There are eggs with about 15 threads present on the one and about 30 on the other pole. As all epithe- mata are already unrolled in the larva, the number of ter- minal knobs per epithema cannot exactly be determined (12-25). The knobs show apical cap-like structures; from each knob 2-4 threads are leading off (Fig. 5 right) (for
MALZACHER, TWO NEW CAENIS FROM NORTH-EASTERN CHINA 65
detailed description of coiled rope-like epithema types see MALZACHER 2011: 73, fig. 13). Micropyle with a large oval sperm guide; mouth broadly rounded; visible part of micropyle channel very short (Fig. 5 left).
Larva
Measurements and colouration
Male larva of last instar, body length 5.7mm, length of cerci 4.5mm; female larva of last instar, body length 8.0 mm, length of cerci 6.0 mm.
Colouration and pigmentation of the ULMER material is not preserved.
Morphology
Cuticle with small granules (pro and mesonotum) or denticles (operculate gills), two types of bristles: (1) very long and thin, (2) very short, broadly spatulate, more or less frayed. The latter form one or two dense rows later- ally and sublaterally on wing buds.
Head: Genae not bulging out. Dorsolateral field of moderate or long bristles on mandibles. Second segment of labial palp 1.2-1.4 times as long as the third (along the centre-line); outer margin with about 6 very long, strong bristles. Sides of postmentum posteriorly slightly diverg- ing, with strong and pointed bristles.
Thorax: Sides of pronotum straight and more or less parallel, anterior corners rounded. Sides of mesonotum with an anterolateral, inconspicuous, flat bulge. Coxal processes narrow, sickle-shaped, not bulging out. Fore femur with a longitudinal row of long thin bristles on dor- sal side. Fore tarsus ventrally with a row of strong, dagger- shaped bristles. Mid tarsus with a similar row, additionally with a few apical pinnate bristles. Hind tarsus ventrally with two rows of dagger-shaped bristles, apical part of the inner row with pinnate bristles. Claws with 7-10 strong denticles. All claws relatively small.
Abdomen: Abdominal segments with long posterolat- eral processes, strongly protruding laterally, particularly in
female larvae; margins with long bent bristles, shorter on posterior segments (Fig. 1b-e). Posteriomedian process of tergum II equilateral triangular, with pointed tip; tergum I also with a short and rounded process. Hind margin of ter- gum VII with long bristles, tergum VIII with only a few bristles (male) or bristles lacking (female), hind margin of terga IX and X with denticles, tergum IX only with a few lateral ones. Hind part of sternum IX triangular, posteriorly broadly rounded or cut, with bristles of moderate length more or less bent medially (Fig. 1b); dorsal side without sha- green. Margins of operculate gills densely provided with long thin bristles (Fig. la). Y-shaped ridges well developed, with a few short basal bristles. A ventral band of microtri- chia consisting of short transverse rows of about 4 micro- trichia (Fig. 6); the band nearly reaching the posteromedian corner of the gill (Fig. la). Microtrichia slightly elongated in the posterolateral part of the row (Fig. 6 left). A couple of spatulate, shortly frayed bristles basally on ventral side. Gill II 2.5 times length of gill I. Gills III-V(V]) with long mar- ginal filaments; a great part of them with 5 or 6 branches.
Differential diagnosis
Caenis wui n.sp. can be distinguished from all other Caenis species by the following combination of characters:
Male imago: Lateral processes on abdomen long or very long. Penis anvil-shaped. Forcipes moderately short with converging sides and a sclerotized apical knob.
Eggs: with two coiled rope epithemata of robusta sub- type. About 15-30 threads are anchored on the egg pole. Chorion granulated, without network of ridges. Micropyle channel very short.
Larva: Second segment of labial palp about 1.25 times as long as the third one. Sides of pronotum straight and parallel. Abdominal segments with long posterolateral processes, laterally protruding. Hind part of sternum IX triangular, dorsal side without shagreen. Operculate gill with a ventral band of microtrichia consisting of short transverse rows each with about 4 microtrichia. Most fila- ments of gills III—V with 5 or 6 branches.
Tab. 1. Differential diagnostic characters of Caenis wui n. sp. and C. robusta.
Sides of pronotum straight, parallel, posterior corner rounded anterior fourth abruptly bent laterally, anterior corner pointed
Operculate gill, band of microtrichia
short transverse rows with about 4 microtrichia | longer transverse rows with 6-8 microtrichia
Hind margin of tergum VII without denticles, with a few bristles or with denticles and a few bristles without bristles
Shagreen on sternum IX without shagreen evenly covered with small denticles
66 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
Caenis wui is closely related to the Palaearctic species Caenis robusta Eaton, 1884 (compare description and fig- ures in MALZACHER 1984 and 1986). Both species share the following characters: Imago: Abdomen with very long lat- eral filaments. Penis anvil-shaped. Forcipes short, with sides converging to the apex. Eggs with coiled rope-like epithemata of robusta subtype (MALZACHER 1982, 2011). — Larva: Third segment of labial palp very long. Fore femur with a longitudinal row of bristles (nearly all other spe- cies with a transverse row). Ventral side of operculate gill with a band of microtrichia consisting of short transverse rows; except for Caenis horaria (Linnaeus, 1758) all other known larvae of Caenis with a single row of microtrichia. The species of the Brachycercinae tribe Caenoculini how- ever possess bands of short transverse rows (MALZACHER, in preparation). For characters distinguishing Caenis wui and C. robusta see Tab. 1.
Zuou et al. (2000) described a species with similar gen- italia, Brachycercus parviforcipes, that was subsequently transferred to the genus Caenis (ZHOU & ZHENG 2004). This species also seems to be similar to Caenis wui, but
Neue Serie 9
the forcipes of the former are short, triangular and pointed, roughly like in Caenis robusta. Unfortunately larvae, and hence arrangement of microtrichia on operculate gill and other larval diagnostic features, are unknown.
3.2. Caenis pekinensis n. sp. (Pigs.2:.7)
Holotype, ¢@ larva (on microslide): M11, China, Peking [= Beijing env.], [date missing], leg. C. F. Wu.
Paraty pes: same data as holotype, 2 9° larvae (differen- tial diagnostic characters are the same in male and female larvae).
Etymology The species epithet refers to the locus typicus Peking [= Bejing].
Male imago
The holotype is a last instar male larva. Therefore a few subimaginal characters can be described:
Base of antennal flagellum not dilated. Abdomen with short posterolateral processes (Fig. 2c). Sternite IX and
=m ae“ . eer ae
Fig. 2. Caenis pekinensis n. sp., larva (a—e) [genitalia of 4 subimago (b) and lateral processes of subimago (c) visible], male subimago (f). —a. Operculate gill, general view. b. Sternum IX. c. Abdominal segment VII, lateral part. d. Lateral outline of pronotum and ante- rior part of mesonotum. e. Coxal processes of mid (right) and hind leg (left). f. Forceps.
MALZACHER, TWO NEW CAENIS FROM NORTH-EASTERN CHINA 67
Figs. 3-7. Caenis wui n. sp. (3-6) [eggs dissected from 9 last instar larva], C. pekinensis n. sp. (7). —3. Egg. 4. Egg, polar region, with tuft of threads. 5. Egg, threads with terminal knobs (right), micropyle (left). 6. Larva, operculate gill, microtrichia from the band on ventral side, from anterolateral (right) and posterolateral (left) regions. 7. Larva, operculate gill, microtrichia from the band on ven- tral side.
68 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
subimaginal genitalia as in Fig. 2b. Penis square, no lat- eral lobes. Forcipes long and straight, median part with nearly parallel sides; apex with a few short spines (Fig. 2f).
Larva
Measurements and colouration
Male larva of last instar, body length 3.5mm, length of cerci 2.5mm; female larva of last instar, body length 6.0mm, length of cerci 3.5 mm.
Colouration and pigmentation of the ULMER material is not preserved.
Morphology
Cuticle smooth or with very small, inconspicuous den- ticles.
Head: Genae bulging out. Dorsolateral field of long bristles on mandibles. Second segment of labial palp 2.2— 2.5 times as long as the third (along the centre-line); outer margin with about 9 long, strong bristles.
Thorax: Sides of pronotum straight, more or less diverging anteriorly, anterior corners rounded; sides of mesonotum anterolaterally with a flat, broadly rounded process (Fig. 2d). Coxal processes well developed, semi- circular or tongue-shaped, margin denticulated (Fig. 2e). Fore femur dorsally with a transverse row of thin bristles of different length and a posteriorly adjoining group of bristles. Mid and hind femora with long bristles on mar- gins and dorsal surface. Tibiae with a submarginal row of long bristles and spines on anterior margin; posterior mar- gin with bristles shorter than those on femora. Fore tar- sus ventrally with a row of short pointed bristles, apical ones more or less pinnate. Mid tarsus with a similar row, additionally with a few apical pinnate bristles. Hind tar- sus ventrally with one long and one short row (short one half the length of long one) or an irregular band of short, pointed bristles with apical ones pinnate. Claws slightly bowed, with minute denticles, often hardly visible.
Abdomen: Abdominal segments with moderate poste- rolateral processes, more or less protruding laterally; mar- gins with long bent bristles, shorter on posterior segments (Fig. 2b, c). Posteriomedian process of tergum II flat and broadly rounded, semicircular. Hind margin of terga VII and VIII with long bristles, of terga IX and X with den- ticles. Hind margin of sternum IX with a flat indenta- tion or cut, with bristles of moderate length more or less bent medially, the median ones apically bifurcated; dor- sal side with one or two rows of microdenticles close to the hind margin (in the female larva immediately on hind margin and hardly visible) (Fig. 2b). Margins of opercu- late gills densely provided with thin bristles of moderate length; surface with scattered thin bristles; posterior part of median ridge reduced, only indicated by a row of thin bristles (Fig. 2a). Outer ridge consisting of strong, rounded
Neue Serie 9
denticles. Ventral row of short microtrichia (Fig. 7) run- ning to the posteriomedian corner of the gill (Fig. 2a). Basal part with more or less frayed, spatulate bristles of different length and shape. Gill IT 2.5 times length of gill I.
Differential diagnosis
Caenis pekinensis n.sp. can be distinguished from all other Caenis species by the following combination of characters:
Male imago: Penis square. Forcipes long and straight, nearly parallel-sided, with a few short apical spines.
Larva: Second segment of labial palp 2.2—2.5 times length of third segment. Sides of pronotum diverging anteriorly. Coxal processes well developed. Posterome- dian process on abdominal tergum II broad, semicircu- lar. Hind margin of sternum IX with a flat indentation or cut. Y-shaped ridge on operculate gill posteriorly reduced, outer ridge consisting of strong rounded denticles.
As a Palaearctic species, Caenis pekinensis seems to belong to the Caenis beskidensis-group. The larva of Cae- nis pycnacantha from eastern China, described by Jia et al. 2010, is similar to that of C. pekinensis, but C. pycna- cantha differs by sides of pronotum clearly convex, claws with 6-10 well developed denticles, and posterior part of Y-shaped ridge well developed. Males of C. pycnacan- tha show a dilated base of antennal flagellum, penis with broadly rounded lobes, bowed forceps with converging sides and strong apical spines.
4 References
Jia, Y.-Y., Qin, J.-Z., Ju, M. & Zuou, C.-F. (2010): A new mayfly spe- cies of Caenis from headwater of Zijin Hill (Nanjing, Eastern China) (Ephemeroptera: Caenidae). — Zootaxa 2535: 61-68.
MALZACHER, P. (1982): Eistrukturen europäischer Caeni- dae (Insecta, Ephemeroptera). — Stuttgarter Beiträge zur Naturkunde, Serie A (Biologie) 356: 15 pp.
MALZACHER, P. (1984): Die europäischen Arten der Gattung Caenis Stephens (Ephemeroptera, Caenidae). — Stuttgarter Beiträge zur Naturkunde, Serie A (Biologie) 373: 48 pp.
MALZACHER, P. (1986): Diagnostik, Verbreitung und Biologie der europäischen Caenis-Arten (Ephemeroptera: Caenidae). — Stutt- garter Beiträge zur Naturkunde, Serie A (Biologie) 387: 41 pp.
MALZACHER, P. (2011): The West African species of Caenis Ste- phens (Insecta: Ephemeroptera) — Stuttgarter Beiträge zur Naturkunde A, Neue Serie 4: 43-74.
ULMER, G. (1936): Neue chinesische Ephemeropteren, nebst Übersicht über die bisher aus China bekannten Arten. — Peking Natural History Bulletin 10: 201-215.
ZHoU, C.-F. & ZHENG, L. (2004): A preliminary study on the genus Caenis (Ephemeroptera: Caenidae) from Chinese mainland, with description of a new Species. — Entomotax- onomia 26: 1-7.
ZHou, C.-F., Gut, H. & Su, C.-R. (2000): The first record of the genus Brachycercus in China with descripton of a new spe- cies (Ephemeroptera: Caenidae). — Entomologia Sinica 7: 132-134.
MALZACHER, TWO NEW CAENIS FROM NORTH-EASTERN CHINA
Author’s address:
Dr. PETER MALZACHER, Friedrich-Ebert-Straße 63, 71638 Ludwigsburg, Germany; e-mail: malzacher.Ib@t-online.de
Manuscript received: 27.V.2015, accepted: 3.1X.2015.
69
BHL in An
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Stuttgarter Beiträge zur Naturkunde A, Neue Serie 9: 71—85; Stuttgart, 30.1V.2016. DOI: 10.18476/sbna.v9.a6 71
New tachyine species from the Oriental Region (Coleoptera: Carabidae: Bembidiini: Tachyına)
MARTIN BAEHR
Abstract
Eight new species of the bembidiine subtribe Tachyina are described from various countries in the Oriental Region: genus Elaphropus Motschulsky, 1839, subgenus Tachyura Motschulsky, 1862: E. serrulipennis n.sp. and E. schawalleri n. sp. from the island of Leyte/Philippines, E. rufinus n. sp. from Brunei/Borneo, E. grimmi n. sp. from Sabah/Borneo, and E. martensi n.sp. from Nepal; genus Tachyta Kirby, 1837: T. quadrinotata n.sp. from Nepal and 7: /aticollis n. sp. from Sabah/Borneo; genus Paratachys Casey, 1918: P. leytensis n. sp. from the island of Leyte/Philippines. For 7: /aticollis n. sp. a new subgenus Eurytachyta within the genus Tachyta is described. Male and female genitalia are figured if available. The new species are compared with their most similar relatives.
Key words: Coleoptera, Carabidae, Bembidiini, Tachyina, new species, Oriental Region.
Zusammenfassung
Acht neue Arten der Subtribus Tachyina der Tribus Bembidiini aus verschiedenen Landern der Orientali- schen Region werden beschrieben: Gattung Elaphropus Motschulsky, 1839, Untergattung Zachyura Motschulsky, 1862: E. serrulipennis n. sp. und E. schawalleri n. sp. von der Philippineninsel Leyte, E. rufinus n. sp. aus Brunei/ Borneo, E. grimmi n. sp. aus Sabah/Borneo und E. martensi n. sp. aus Nepal; Gattung Tachyta Kirby, 1837: 7: qua- drinotata n. sp. aus Nepal und T: Jaticollis n.sp. aus Sabah/Borneo; Gattung Paratachys Casey, 1918: P. leytensis n.sp. von Leyte. Fur Tachyta laticollis n.sp. wird eine neue Untergattung Eurytachyta innerhalb der Gattung Tachyta beschrieben. Die männlichen und weiblichen Sexualorgane werden abgebildet, sofern bekannt. Die neuen Arten werden mit ähnlichen bzw. verwandten Arten verglichen.
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1 Introduction
Through courtesy of several collectors and museum curators a number of tachyine species from the Oriental Region have been received recently for identification that include some undescribed species alongside with several more or less well known species. The new species belong to the genera Elaphropus Motschulsky, 1839, subgenus Tachyura Motschulsky, 1862, Tachyta Kirby, 1837, and Paratachys Casey, 1918, and are described herein.
The supraspecific taxonomy of the subtribe (or tribe) Tachyina (-in1) is still being discussed and several opin- ions exist about the status of most supraspecific groups. In this paper I mainly follow the system of Lorenz (2005) who, apart from a number of well defined genera, for the bulk of species acknowledged two genera, namely Tachys Dejean, 1821 and Elaphropus Motschulsky, 1839. Tachy- ura Motschulsky, 1862 belongs as subgenus to Elaphropus.
The subgenus Tachyura as denoted by Lorenz (2005) (including the Nearctic species of the former genus Bary- tachys Chaudoir, 1868) covers more than 250 species that are distributed in the Palaearctic, Afrotropical, Oriental, Papuan/Australian, and Nearctic Regions. The subgenus, or at least the eastern species, excluding the Nearctic Bary- tachys, is equivalent to the politus-group and some related species-groups of ANDREWES (1925) and DARLINGTON (1962). It should be mentioned, however, that the generic and subgeneric systematics of Tachyini are by no means settled. Scıaky & VIGNA TAGLIANTI (2003) and Kopecky (2003), for example, proposed different systems with more restricted limits of the subgenus Tachyura. Also the status of the genus or subgenus Nototachys Alluaud, 1930 is con- troversely discussed.
The species of Tachyura in the sense of LorENz (2005) are characterized by absence of deep pits on the mentum, parallel frontal sulci which do not completely cross over
72 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
onto the clypeus, presence of the 8" elytral stria which usually is deeply impressed and almost complete, the api- cal recurrent striole being situated more or less ın the mid- dle of the elytra, edentate humerus, and the pronotum with an impressed basal sulcus.
This subgenus includes very differently shaped, struc- tured, and coloured species. The Oriental region is partic- ularly rich in species and certainly the Oriental Tachyura fauna would merit a modern comprehensive revision, in spite of the large paper of ANDREwEs (1925) which still must be used for identification of species. However, this is pres- ently outside of the capacity of the author. Also probably many additional species are supposed to exist in the Oriental Region which has not been sufficiently and systematically collected for tachyine beetles. The New Guinean species of Tachyura were revised by DARLINGTON (1962), but this revi- sion likewise is rather preliminary. The Australian species were revised by BAEHR (1988). A number of additional spe- cies, mainly from the southern Oriental Region and from New Guinea, were recently described by BAEHR (2014a).
Kopecky (2003) and Lorenz (2005) included Paratachys Casey, 1918 as a subgenus in the genus Tachys Dejean, 1821 (s.1.). In this decision I do not follow them in the present paper, but, like most other modern authors, treat Paratachys as a separate genus. It is a very large genus which includes a multitude of species throughout the world, and it is congruent with the triangularis-group of ANDREWES (1925).
Species of Paratachys are characterized by the presence of two deep pits in the mentum and a distinct recurrent stria on the elytra that runs in the middle of the apex and embraces the posterior discal puncture just at ıts incurved apex.
The genus Tachyta Kirby, 1837 includes a number of more or less depressed species which are distributed in the Holarctic, Afrotropical, Oriental, and Australian Regions. It is characterized by finely serrulate tarsal claws, elon- gate terminal palpomeres, and a straight, elongate recur- rent stria that runs close to the elytral margin. The genus includes three subgenera, two of which occur ın the Ori- ental Region.
Eight new species from the genera Elaphropus, Tachyta, and Paratachys are herein described from Nepal, Borneo, and the Philippines. In the genus Tachyta a new subgenus Eurytachyta 1s described for an unusual new species.
Acknowledgements
I am indebted to ROLAND GRIMM (Neuenbürg), WOLFGANG SCHAWALLER (Stuttgart), and Davip WRraseE (Berlin) for the kind loan or gift of specimens. I also thank CHARLES Huser (Bern) and JoAcHIM Scumipt (Admannshagen) for some corrections.
2 Material and methods
For the taxonomic treatment standard methods were used. The genitalia were removed from specimens relaxed for a night
Neue Serie 9
in a jar under moist atmosphere, then cleaned for a short while in hot 4% KOH.
The habitus photographs were obtained with a digital cam- era using ProgRes Capture Basic and AutoMontage and subse- quently were worked with Corel Photo Paint X14.
Measurements were taken using a stereo microscope with an ocular micrometer. Body length has been measured from apex of labrum to apex of elytra. Length of pronotum was measured along midline, width of base of pronotum at the extreme tips of the basal angles.
The types are stored in the working collection M. BAEHR in Zoologische Staatssammlung, München (CBM), in Staatliches Museum für Naturkunde, Stuttgart (SMNS), and in the collec- tion D. Wrasse, Berlin (CWB).
3 Genus Elaphropus, subgenus Tachyura Motschulsky, 1862
Tachyura Motscuu sky, 1862: 27. — ANDREWES 1925: 328; Csıkı 1928: 166; BAEHR 1988: 229, 2014a: 2; Sctaky & VIGNA TAGLIANTI 2003: 93; Kopecky 2003: 278; LoRENz 2005: 208.
Type species: Elaphrus quadrisignatus Duftschmid, 1812, by subsequent designation by MorscHuLsky (1862).
Diagnosis The subgenus is characterized by absence of deep mental pits, well developed posterior transverse sulcus of the pronotum, almost complete 8" elytral stria, rounded humerus, well developed recurrent stria on the elytra that is situated more or less in the middle, and presence of two discal elytral punctures and setae.
Description of Elaphropus serrulipennis n. sp. (Figs. 1, 6, 12)
Holoty pe (6): “Philippines, Leyte, visca N Baybay, cul- tiv. land, 1991, leg. SCHAWALLER & al. / 4.111.1991” (SMNS).
Paratypes (346, 829): same locality, but 28.11.1991, 1.111.1991, 4.111.1991, 8. 111.1991 (CBM, SMNS).
Etymology
The species name refers to the strongly serrate lateral mar- gins of the elytra.
Diagnosis
Characterized by pale reddish-brown colour, depressed eye, cordiform pronotum with rectangular basal angle, markedly serrate margin of the elytra, presence of the complete set of elytral striae, uninterrupted 8" elytral stria, straight, and oblique recurrent stria that is rather closely situated to the lateral margin.
In some character states this species reminds £. singu- laris (Andrewes, 1925) of the exaratus-group of ANDREWES (1925) or the singularis-group in the sense of DARLINGTON (1962), but is distinguished by the completely striate elytra
BAEHR, NEW TACHYINE SPECIES FROM THE ORIENTAL REGION 13
which are serrulate and setulose almost along their whole length, and by the uninterrupted 8" stria.. Because it does not match the most important determining character states of both groups, I prefer to include it into the subgenus Tachyura without attributing to a definitive subgroup.
Description
Measurements and ratios. Body length: 2.10-2.25 mm; width: 0.85—0.90 mm. Ratios: Width/length of pronotum: 1.42-1.47; widest diameter/width of base of pronotum: 1.23-1.28; width of pronotum/width of head: 1.31-1.35; length/width of elytra: 1.51—1.57.
Colour (Fig. 12). Upper and lower surfaces unicolour- ous pale reddish-brown. Antenna, palpi, and legs yellow.
Head (Fig. 12). Of normal size but eye depressed and little produced laterad, orbit small and indistinct. Labrum anteriorly straight. Clypeus straight, clypeal suture shal- low. Frons slightly convex, without any impression. Fron- tal furrows wide and shallow, slightly curved, anteriorly not crossing onto clypeus, posteriorly attaining middle of eye. Mandible rather elongate, palpi thick, pubescent, api- cal palpomere short and very thin. Mentum without pores. Antenna fairly elongate, 7-10" antennomeres twice as long as wide. Dorsal surface impunctate, with distinct though superficial, about isodiametric microreticulation that consists of rather large meshes, surface glossy,
Pronotum (Fig. 12). Moderately wide, cordiform, at base about as wide as at apex; dorsal surface gently con- vex. Apex with deep excision, apical angles markedly pro- jected and rather acute. Lateral border evenly convex but in basal quarter straight or very slightly concave. Base lat- erally slightly excised, in middle produced, basal angles rectangular, acute. Lateral sulcus very narrow, not wid- ened at base. Both, apex and base not margined. Median line fine, slightly impressed, not attaining the apex, near base deepened to a wide sulcus. Anterior transverse impression barely perceptible, posterior impression deep, finely crenulate, in the median sulcus with a pore. Basal grooves absent, base near lateral margin without a carina. Anterior lateral seta inserted at anterior third, in front of widest diameter, posterior lateral seta inserted near basal angle. Surface impunctate, with extremely superficial, about isodiametric microreticulation, very glossy.
Elytra (Fig. 12). Moderately elongate, lateral mar- gin slightly convex, elytra wıde at humerus; dorsal sur- face moderately convex but depressed on disk. Humerus rounded, not produced, basal margin incomplete, attain- ing c. level of 4" stria. The complete lateral margin ser- rulate, more coarsely so in basal third, but not setulose. All striae distinct, deeply impressed, at bottom very finely crenulate. Striae almost attaining base, except the sutural one ending in front of apex, the outer ones more short- ened than the inner ones, apex widely glabrous. All inter- vals on disk markedly raised, even slightly tectiform. 8"
stria complete, uninterrupted, deeply impressed, impunc- tate. Recurrent stria deep, moderately elongate, oblique, straight, at end not incurved, situated rather close to mar- gin. 3" interval bipunctate, the anterior puncture located at basal third, the posterior puncture located about at apical third, both adjacent to 3" stria. Intervals impunctate, with distinct, though superficial, about isodiametric microre- ticulation, surface glossy.
Lower surface. Not pilose, glabrous. Metepisternum slightly > 1.5 times as long as wide at apex. Terminal abdom- inal sternum in male bisetose, in female quadrisetose.
Legs. Two basal tarsomeres of the male protarsus slightly widened, squamose underneath, and mediad tri- angularly dentate.
Male genitalia (Fig. 1). Aedeagus short and compact, stout in apical part, lower surface evenly concave. Apex short and stout, rounded; internal sac with several slightly twisted folds, one of which is rather heavily sclerotized. Parameres very dissimilar, the left one very large, tri- angular and slightly curved down, the right one smaller but comparatively wide, both with two elongate and one shorter apical setae.
Female gonocoxites (Fig. 6). Gonocoxite 1 compar- atively large, without any setae. Gonocoxite 2 short and wide, slightly triangular, straight, only near apex slightly incurved, with wide, obtuse apex. Two short but stout ven- tro-lateral ensiform setae situated in apical half, one short and stout dorso-median ensiform seta situated close to apex; one short subapical nematiform seta near apex orig- inating from a circular pit.
Variation. Very little variation noted.
Distribution
Island of Leyte, Philippines. Known only from type locality.
Collecting circumstances
According to information of the collector, sampled at light.
Relationships
Uncertain. The species combines character states of different species groups, respectively subgenera, there- fore its status remains uncertain as long as a comprehen- sive phylogenetic survey of the whole tachyine lineage is lacking.
Description of Elaphropus schawalleri n. sp. (Figsa2- 713}
Holotype (&): “Philippines, Leyte, SW Abuyog, 28.11.1991, river bank, leg. SCHAWALLER & al.” (SMNS). Paratypes (444, 4 29): same data (CBM, SMNS).
74 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
Etymology
The species name is a patronym in honour of the collector WOLFGANG SCHAWALLER of Staatliches Museum für Naturkunde, Stuttgart.
Diagnosıs Characterized by presence of three elytral striae, uni- maculate elytra, cordiform pronotum, and dark antenna. Very similar to E. borneensis (Andrewes, 1925), but dis- tinguished by lesser number of elytral striae, narrower base of the pronotum, shorter elytra, and darker antenna, palpi, and legs.
Description
Measurements and ratios. Body length: 2.50-2.75 mm; width: 1.1-1.2mm. Ratios: Width/length of pronotum: 1.35—1.42; widest diameter/width of base of pronotum: 1.26-1.28, width of pronotum/width of head: 1.30-1.33; length/width of elytra: 1.41-1.46.
Colour (Fig. 13). Black, elytra near apex with an ill delimited, rather inconspicuous, yellow or pale red spot that extends between 3” or 4" and 7" intervals. Antenna dark with I and basal part of 2" antennomere pale, basal and apical palpomeres of maxillary palpus dirty yellow, penultimate palpomere dark; legs dirty yellow to pale brown.
Head (Fig. 13). Of average size. Eye large, laterad rather produced, orbit short, oblique-convex. Labrum anteriorly straight. Clypeus straight, clypeal suture incon- spicuous or even not perceptible. Frons slightly convex, without any impression. Frontal furrows slightly dupli- cated, deep, short, straight, anteriorly slightly crossing to the clypeus, posteriorly not turned laterad. Sulcus medi- ally of the eye deep. Mandible, palpi and the lower surface of the head as in similar species of the subgenus. Antenna moderately short, 7-10" antennomeres slightly less than twice as long as wide. Dorsal surface impunctate, with distinct though slightly superficial, about isodiametric microreticulation, fairly glossy.
Pronotum (Fig. 13). Moderately wide, at base barely wider than at apex; dorsal surface gently convex. Apex almost straight, apical angles not projected and widely rounded. Lateral border convex over most of its length, near base shortly concave. Base laterally straight to faintly oblique, in middle slightly produced, basal angles rectan- gular, acute. Lateral sulcus moderately narrow, not wid- ened basad. Apex laterally finely margined, base not margined except near basal angles. Median line fine, slightly deepened basad, neither attaining apex nor base. Anterior transverse impression barely perceptible, poste- rior impression deep, distinctly crenulate, in middle with a small, barely perceptible puncture. Basal grooves moder- ately deep, base near lateral margin with a short, slightly oblique, sharp carina. Anterior lateral seta inserted slightly
Neue Serie 9
in front of middle, about at widest diameter, posterior lat- eral seta inserted at basal angle. Surface impunctate, with indistinct, very superficial, isodiametric to slightly trans- verse microreticulation that in some areas is almost invisi- ble; surface glossy, except for the rather rugose base.
Elytra (Fig. 13). Moderately elongate, not oviform, lat- eral margin in middle almost straight, wide at humerus; dorsal surface moderately convex but depressed on disk. Humerus rounded, basal margin incomplete, attaining c. level of 4" stria. Three deeply impressed, not crenulate striae present. 1 stria attaining scutellary puncture and apex, 2"! and 3" striae at base shortened, and gradually ending far in front of apex, apex widely glabrous. In some specimens in middle faint traces of a 4" and even a 5" stria perceptible. Intervals on disk slightly raised. 8" stria com- plete, deeply impressed, impunctate. Recurrent stria deep, elongate, straight, at end not incurved. 3" interval bipunc- tate, the anterior puncture located about at basal third, the posterior puncture located slightly in front of the api- cal third, both adjacent to 3" stria. Surface impunctate, at least laterally with traces of very superficial, about isodia- metric microreticulation, surface glossy.
Lower surface. Not pilose, glabrous. Metepisternum c. 1.75 times as long as wide at apex. Terminal abdominal sternum in male bisetose, in female quadrisetose.
Legs. Two basal tarsomeres of the male protarsus slightly widened, squamose underneath, and mediad trı- angularly dentate.
Male genitalia (Fig. 2). Aedeagus rather elongate, stout in middle, triangularly narrowed towards apex, lower surface evenly concave. Apex rather elongate, obtusely rounded; internal sac with several slightly twisted folds, one of which, situated at bottom, is heavily sclerotized. Parameres very dissimilar, the left one very large, trian- gular, with two elongate apical setae, the right one smaller but comparatively wide, with a single elongate apical seta.
Female gonocoxites (Fig. 7). Gonocoxite 1 compar- atively large and elongate, triangular, without any setae. Gonocoxite 2 elongate, triangular, curved, with rather acute apex. Two very small ventro-lateral ensiform setae situated about in middle, moderately separated, the dorso- median ensiform seta absent; one short subapical nemati- form seta present near apex and originating from a circular
pit. Variation. Very little variation noted.
Distribution
Island of Leyte, Philippines. Known only from type locality.
Collecting circumstances
Sampled at “river bank”, but unknown whether by hand collecting or whether at light.
BAEHR, NEW TACHYINE SPECIES FROM THE ORIENTAL REGION 75
Relationships
In body shape, structure of elytra, and aedeagus very similar, and probably closely related to E. borneensis (Andrewes, 1925).
Description of Elaphropus rufinus n. sp. (Figs. 3, 14)
Holoty pe (2): “Borneo, Brunei, Temburong Kuala Bela- long, BoRCHERDING leg. / 10.11.1995 HW KBFSC” (CWB). Paratype (d): same data (CBM).
Etymology The name refers to the dark rufous colour of this species.
Diagnosis
Characterized by rufous colour, cordiform prono- tum with trifoveate basis, presence of four short, coarsely crenulate elytral striae, inconspicuously quadrimaculate elytra, and absolutely glabrous surface. Distinguished from similarly short and compact species [e.g. E. latus (Peyron, 1858) or E. aeneus (Putzeys, 1875)] by colour and number, size, and structure of the elytral striae.
Description
Measurements and ratios. Body length: 2.55-2.70 mm; width: 1.25-1.30 mm. Ratios: Width/length of pronotum: 1.27-1.30, widest diameter/width of base of pronotum: 1.27-1.28, width of pronotum/width of head: 1.26-1.27, length/width of elytra: 1.35—1.37.
Colour (Fig. 14). Upper and lower surfaces dark red, elytra with four moderately distinct pale red spots. The humeral spot more ill delimited and extended from 4" or 5“ interval to margin, the apical spot more distinct, extended from 3™ to 7" intervals, and well removed from apex that is slightly paler than the disk. 1-3" antenno- meres and basal half of 4" yellow, rest infuscate. Palpi and legs yellow.
Head (Fig. 14). Of average size. Eye very large, lat- erad well produced, orbit very small. Labrum anteriorly slightly excised. Clypeus straight, clypeal suture rather deep. Frons slightly convex, without any impression. Frontal furrows duplicated, deep, almost straight, anteri- orly slightly crossing to the clypeus, posteriorly slightly turned laterad. Sulcus medially of the eye deep. Mandi- ble, palpi and the lower surface of the head as in simi- lar species of the subgenus. Antenna moderately elongate, 7"_10" antennomeres c. 1.75 times as long as wide. Dorsal surface impunctate and without microreticulation, very glossy, only labrum with superficial, isodiametric micro- reticulation.
Pronotum (Fig. 14). Rather wide, at base slightly wider than at apex; dorsal surface rather convex. Apex straight,
apical angles not projected, very widely rounded. Lateral border convex, in basal third rather deeply concave. Base laterally straight, in middle produced, basal angles slightly less than rectangular, acute. Lateral sulcus narrow, not widened basad. Apex laterally finely margined, base not margined except near basal angles. Median line very fine, inconspicuous, neither attaining apex nor base. Anterior transverse impression not perceptible, posterior impres- sion deep, very finely crenulate, in middle with three large and deep pores. Basal grooves very deep, base near lateral margin with a short, slightly oblique, sharp carina. Anterior lateral seta inserted slightly in front of middle, at widest diameter, posterior lateral seta inserted at basal angle. Sur- face very glossy, impunctate and without microreticulation.
Elytra (Fig. 14). Wide and short, not oviform, lateral margin in middle straight, wide at humerus; dorsal sur- face convex but depressed on disk. Humerus faintly angu- late, basal margin incomplete, attaining c. level of 4" stria. Four striae present, deeply impressed, and coarsely cren- ulate. 1° stria attaining scutellary puncture and apex, 2™_4" striae gradually shortened at base and apex, apex widely glabrous. All intervals on disk raised. 8" stria com- plete, deeply impressed, impunctate. Recurrent stria fairly deep, short, oblique, at end incurved. 3™ interval bipunc- tate, the anterior puncture located at basal third, the pos- terior puncture located slightly in front of the apical third, both adjacent to 3" stria. Intervals impunctate and without microreticulation, surface very glossy.
Lower surface. Not pilose, glabrous. Metepisternum slightly <1.5times as long as wide at apex. Terminal abdominal sternum in male bisetose.
Legs. Two basal tarsomeres of the male protarsus slightly widened, squamose underneath, and mediad trı- angularly dentate.
Male genitalia (Fig. 3). Aedeagus moderately elongate, stout in middle, triangularly narrowed towards apex, lower surface straight. Apex rather short, fairly stout, obtusely rounded; internal sac with several slightly twisted folds, one of which is heavily sclerotized. Parameres very dis- similar, the left one very large, triangular, the right one smaller but comparatively wide, both with three elongate apical setae.
Female gonocoxites. Unknown.
Variation. Very little variation noted.
Distribution Brunei, Borneo. Known only from type locality.
Collecting circumstances Not recorded.
Relationships
A rather unique species, apparently without close rel- atives.
76 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
Description of Elaphropus martensi n. sp. (Figs. 4, 8, 15)
Holotype (8): “320 Ilam Distr. betw. Mai Pokhari and Gitang Khola Valley, 2100-1750 m, tree rich cultural land, 11 Apr 1988, J. MARTENS & W. SCHAWALLER leg. / Nepal-Expedi- tionen JOCHEN MARTENS” (SMNS).
Paratype (9): “321 Iam Distr., Gitang Khola Valley, Alnus forest along river, 1750 m, 11-13 Apr 1988, J. MARTENS & W. SCHAWALLER leg. / Nepal-Expeditionen JOCHEN MARTENS” (CBM).
Etymology
The name is a patronym in honour of the collector JocHEN MARTENS, well known explorer of the fauna of Nepal.
Diagnosis Characterized by black colour, cordiform pronotum with unifoveate basis, presence of only one distinctly impressed elytral stria, quadrimaculate elytra, and abso- lutely glabrous surface. Distinguished from the most sim- ilar species E. stevensi (Andrewes, 1925) by absence of the 2" stria and distinctly crenulate base of the pronotum.
Description
Measurements and ratios. Body length: 2.85—2.90 mm; width: 1.25mm. Ratios: Width/length of pronotum: 1.33— 1.34; widest diameter/width of base of pronotum: 1.36— 1.40; width of pronotum/width of head: 1.36—1.39; length/ width of elytra: 1.44—1.45.
Colour (Fig. 15). Upper and lower surfaces black, elytra with four moderately distinct pale red spots. The humeral spot slightly triangular and extended from site of 4" interval to margin, the apical spot transverse, extended from 3" interval to margin and well removed from apex that is slightly paler than the disk. Antenna, palpi, and legs yellow.
Head (Fig. 15). Of average size. Eye fairly large, lat- erad moderately produced, orbit comparatively elon- gate, c. % of eye length, oblique-convex. Labrum almost straight. Clypeus straight, clypeal suture distinct. Frons slightly convex, without any impression. Frontal furrows duplicated, short, deep, anteriorly slightly crossing to the clypeus, posteriorly turned laterad. Sulcus medially of the eye deep. Mandible, palpi and the lower surface of the head as in similar species of the subgenus. Antenna moder- ately elongate, 7-10" antennomeres slightly < 1.75 times as long as wide. Dorsal surface impunctate and without microreticulation, very glossy, only labrum with superfi- cial, isodiametric microreticulation.
Pronotum (Fig. 15). Wide, at base slightly wider than at apex; dorsal surface rather convex. Apex straight, api- cal angles not projected, very widely rounded. Lateral bor- der convex, in basal fourth more or less deeply concave. Base laterally straight, in middle produced, basal angles slightly less than rectangular, acute, laterad even slightly projected. Lateral sulcus narrow, slightly widened in mid-
Neue Serie 9
dle, narrowed basad. Apex laterally finely margined, base not margined except near basal angles. Median line fine, slightly deepened, neither attaining apex nor base. Anterior transverse impression not perceptible, posterior impression deep, distinctly crenulate, in middle with a large and deep pore. Basal grooves very deep, base near lateral margin with a short, slightly oblique, sharp carina. Anterior lateral seta inserted slightly in front of middle, slightly in front of widest diameter, posterior lateral seta inserted at basal angle. Surface very glossy, impunctate and without microreticulation.
Elytra (Fig. 15). Moderately elongate, not oviform, lateral margin in middle almost straight, wide at humerus; dorsal surface convex but depressed on disk. Humerus not angu- late, obliquely rounded, basal margin incomplete, attain- ing c. level of 4" stria. Sutural stria distinct and complete, deeply impressed, not or very weakly crenulate; traces of 2™ stria in middle more or less distinct, but barely impressed. 8" stria complete, deeply impressed, impunctate. Recurrent stria fairly deep, short, oblique, at end slightly incurved. 3" interval bipunctate, the anterior puncture located at basal third, the posterior puncture located slightly behind middle, both adjacent to position of 3 stria. Surface impunctate and without microreticulation, very glossy.
Lower surface. Not pilose, glabrous. Metepisternum c. 1.5times as long as wide at apex. Terminal abdominal sternum in male bisetose, in female quadrisetose.
Legs. Two basal tarsomeres of the male protarsus slightly widened, squamose underneath, and mediad trı- angularly dentate.
Male genitalia (Fig.4). Aedeagus rather elongate, moderately stout ın middle, triangularly narrowed towards apex, lower surface in middle slightly convex, but apical fourth markedly curved down. Apex very stout but rather elongate, obtusely rounded; internal sac with several slightly twisted folds, one of which is heavily sclerotized. Parameres very dissimilar, the left one very large, trian- gular, the right one smaller and narrow, both with a single elongate apical seta.
Female gonocoxites (Fig. 8). Gonocoxite 1 large, trian- gular, without any setae. Gonocoxite 2 fairly elongate, tri- angular, slightly curved, with rather acute apex. Two very small ventro-lateral ensiform setae situated about in mid- dle, moderately separated, the dorso-median ensiform seta absent; one short subapical nematiform seta present near apex and originating from a circular pit.
Variation. Little variation noted in depth of sinuosity of the lateral margin of the pronotum and in distinctness of 2" stria.
Distribution
Nepal. Known only from the Singalila mountain range of the easternmost Nepal Himalaya close to the border to the Darjeeling District of India.
BAEHR, NEW TACHYINE SPECIES FROM THE ORIENTAL REGION 77
Collecting circumstances
Both specimens collected in forested area 1n river val- ley, but unknown whether by hand collecting or whether at light.
Relationships
In body shape and structure of elytra most similar and probably closely related to E. stevensi (Andrewes, 1925).
Description of Elaphropus grimmi n. sp. (Figs. 5, 16)
Holoty pe (&): “Borneo, Malaysia, Sabah NE, Keningau, Bingkor, 380 m, 22.111.2013, R. Grimm” (CBM).
Etymology
The species name is a patronym in honour of the collector ROLAND GRIMM, well known authority of Oriental Tenebrioni- dae.
Diagnosıs
Characterızed by unicolourous glossy black colour and hirsute elytra. Distinguished from the likewise hir- sute E. barringtoni (Andrewes, 1925), E. interpunctatus (Putzeys, 1875), and E. hirsutus Baehr, 2014, by different number of elytral striae, in addition from E. interpuncta- tus by absence of the elytral spot, from E. barringtoni by the elytral striae almost attaining the base of the elytra, and from £. hirsutus by not deeply excised base of the pro- notum.
Description
Measurements and ratios. Body length: 2.6 mm; width: 1.1mm. Ratios: Width/length of pronotum: 1.32; widest diameter/width of base of pronotum: 1.34; width of prono- tum/width of head: 1.38; length/width of elytra: 1.47.
Colour (Fig. 16). Upper and lower surfaces unicolour- ous glossy black, elytra unspotted. Antenna dark except 1‘ and basal half of 2” antennomere. Palpi yellow, but the penultimate palpomeres piceous. Legs bright yellow.
Head (Fig. 16). Of average size. Eye large, laterad well produced, orbit very small. Labrum anteriorly straight. Clypeus straight, clypeal suture shallow. Frons slightly convex, without any impression. Frontal furrows not perceptibly duplicated, deep, slightly oblique, anteri- orly slightly crossing to the clypeus, posteriorly slightly turned laterad. Sulcus medially of the eye deep. Mandi- ble, palpi and the lower surface of the head as in simi- lar species of the subgenus. Antenna moderately short, 7"_10" antennomeres slightly < 1.5 times as long as wide. Dorsal surface impunctate and without microreticulation, very glossy, only labrum with superficial, isodiametric microreticulation.
Pronotum (Fig. 16). Moderately wide, at base barely wider than at apex, dorsal surface very convex. Apex straight, apical angles not projected and widely rounded. Lateral border very convex, in basal fourth slightly con- cave. Base slightly convex, laterally slightly oblique, basal angles slightly more than rectangular, acute. Lateral sulcus very narrow, basad not widened. Apex laterally finely mar- gined, base not margined except near basal angles. Median line very fine, inconspicuous, barely impressed, neither attaining apex nor base. Anterior transverse impression barely perceptible, posterior impression deep, little curved, distinctly crenulate, in middle with a very small, barely discernible puncture. Basal grooves deep, base near lateral margin with a very short, slightly oblique, sharp carina. Anterior lateral seta inserted about in middle, at widest diameter, posterior lateral seta inserted at basal angle. Sur- face very glossy, impunctate and without microreticulation.
Elytra (Fig. 16). Fairly elongate, not oviform but some- what oblong-triangular, widest at humerus, lateral mar- gin in middle straight and very slightly oblique; dorsal surface convex but depressed on disk. Humerus oblique and rounded, basal margin incomplete, attaining c. level of 3" stria. Five striae present, deeply impressed, smooth. All striae, including 1‘, at base slightly shortened, the inner ones slightly more than 5" stria. All striae except the sutural one ending far in front of apex, the outer ones more shortened than the inner ones, apex widely glabrous. All intervals on disk markedly convex. 8" stria complete, deeply impressed, impunctate. Recurrent stria short, deep, oblique, at end not incurved. Intervals with a row of elon- gate, erect setae which is much denser on the even inter- vals, surface therefore remarkably hirsute. Punctures on 3 interval not discernible. Surface without microreticu- lation, very glossy.
Lower surface. Not pilose, glabrous. Metepisternum slightly <1.5times as long as wide at apex. Terminal abdominal sternum in male quadrisetose.
Legs. Two basal tarsomeres of the male protarsus barely widened, faintly squamose underneath, and mediad extremely indistinctly triangularly dentate.
Male genitalia (Fig. 5). Aedeagus short and compact, very stout in apical half, lower surface slightly concave. Apex very short and widely rounded; internal sac with several slightly twisted but little sclerotized folds. Para- meres very dissimilar, the left one very large, triangular, the right one smaller, both with three elongate apical setae.
Female gonocoxites. Unknown.
Variation. Unknown.
Distribution Sabah, Borneo. Known only from type locality.
Collecting circumstances Not recorded.
78 STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
Figs. 1-5. Male genitalia, aedeagus left side, right and left parameres. — 1. Elaphropus serrulipennis n. sp. 2. E. schawalleri n. sp. 3. E. rufinus n. sp. 4. E. martensi n. sp. 5. E. grimmi n. sp. — Scale bars: 0.2 mm.
BAEHR, NEW TACHYINE SPECIES FROM THE ORIENTAL REGION 79
Figs. 6-11. Female gonocoxites. — 6. Elaphropus serrulipennis n. sp. 7. E. schawalleri n. sp. 8. E. martensi n. sp. 9. Tachyta (Parata- chyta) quadrinotata n. sp. 10. T: (Eurytachyta) laticollis n. sp. 11. Paratachys leytensis n. sp. — Scale bars: 0.2 mm.
80 STUTTGARTER BEITRAGE ZUR NATURKUNDE A Neue Serie 9
Figs. 12-19. Habitus. — 12. Elaphropus serrulipennis n.sp. 13. E. schawalleri n.sp. 14. E. rufinus n.sp. 15. E. martensi n. sp. 16. E. grimmin. sp. 17. Tachyta (Paratachyta) quadrinotata n. sp. 18. T. (Eurytachyta) laticollis n. sp. 19. Paratachys leytensis n. sp. — Scale bars: 1 mm.
BAEHR, NEW TACHYINE SPECIES FROM THE ORIENTAL REGION 81
Relationships
In body shape and structure of elytra very similar, and probably closely related, to E. barringtoni (Andrewes, 1925) from the Philippines and E. interpunctatus (Putzeys, 1875) from Sulawesi.
4 Genus Tachyta Kirby, 1837
Tachyta Kirpy, 1837: 56. — Csıkı 1928: 166; Erwin 1975: 5; ScIAKY & VIGNA TAGLIANTI 2003: 92; Kopecky 2003: 277; LORENZ 2005: 207; BAEHR 2014b: 8.
Type species: Tachyta picipes Kirby, 1837, by monotypy [= Tachyta nana inornata (Say, 1823)].
Diagnosis
The genus is characterized by absence of deep men- tal pits, elongate terminal palpomeres, finely serrulate tar- sal claws, and a straight, elongate recurrent stria that runs close to the lateral margin of the elytra.
Three subgenera have been described, two of which occur in the Oriental region, namely the nominate sub- genus that covers depressed, mostly strongly microreticu- lated species, and Paratachyta Erwin, 1973 that includes a few small, less depressed, glabrous species. The third sub- genus Australotachyta Baehr, 2013, is only known from northern Australia.
An additional subgenus is herein described for a single species of quite aberrant body shape that, however, pos- sesses all determining character states of the genus.
4.1 Subgenus Paratachyta Erwin, 1975
Paratachyta Erwin, 1975: 7. — SclAaky & VIGNA TAGLIANTI 2003: 92; Kopecky 2003: 277; LoRENz 2005: 207.
Type species: Tachys coracinus Putzeys, 1875, by original designation.
Diagnosis The subgenus originally was characterized by moder- ately convex body shape, glabrous surface, and presence of only the I and 8" elytral striae. In the meantime, however, a few species have been described that match body shape and glabrous surface, but possess more than one discal stria. The new species below likewise belongs to this group.
Description of Tachyta (Paratachyta) quadrinotata n. sp. (Figs. 9, 17)
Holotype (@): “320 Ilam Distr., betw. Hodia Khola Valley to Sktia, 250-500 m, Shorea forest, dry, 7 Apr 1988, MARTENS & SCHAWALLER leg. / Nepal-Expeditionen JOCHEN MARTENS” (SMNS).
Etymology The species name refers to the quadrimaculate elytra.
Diagnosis
Characterized by presence of three elytral striae and traces of additional ones, and glabrous, not microreticu- late, quadrimaculate elytra. Distinguished from the sim- ilarly quadrimaculate species 7. quadriplagiata Baehr, 2014 from Vietnam by lesser body size, shorter elytra, presence of only three, instead of five well developed ely- tral striae, and dark femora.
Description
Measurements and ratios. Body length: 2.25mm; width: 1.05mm. Ratios: Width/length of pronotum: 1.45; widest diameter/width of base of pronotum: 1.11; width of pronotum/width of head: 1.43; length/width of elytra: 1.33.
Colour (Fig. 17). Black, elytra with two pale red spots. The humeral spot extended from 4" interval to margin but not touching the base, the apical spot extended between 34 and 6" interval and far removed from apex. Palpi, antenna, and legs bright yellow, but femora infuscate.
Head (Fig. 17). Of average size. Eye large, laterad well produced, orbit very small. Labrum anteriorly straight. Clypeus straight, clypeal suture barely indicated. Frons convex, without any impression. Frontal furrows elongate, deep, curved, attaining the posterior margin of the eye. Apical palpomeres elongate. Antenna short, 7-10" anten- nomeres slightly longer than wide. Dorsal surface impunc- tate and without microreticulation, very glossy, only labrum with superficial, isodiametric microreticulation.
Pronotum (Fig. 17). Wide, at base about as wide as in middle, narrowed to apex; dorsal surface gently convex. Apex with very shallow excision, apical angles barely pro- jected and very widely rounded. Lateral border in basal half very slightly concave. Base laterally straight, in mid- dle faintly produced, basal angles rectangular. Lateral sul- cus deep, anteriorly narrow, basad slightly widened. Apex laterally finely margined, base laterally margined. Median line deeply sulcate, deepened basad, attaining apex and base. Anterior transverse impression only in parts percep- tible, posterior impression deep, smooth, in middle with an elongate sulcus. Basal grooves barely perceptible, base near lateral margin with a short, slightly oblique carina. Anterior lateral seta inserted at apical third, posterior lat- eral seta inserted at basal angle. Surface impunctate, with- out microreticulation, very glossy.
Elytra (Fig. 17). Wide and short, oviform, lateral mar- gin even in middle slightly convex; dorsal surface mod- erately convex. Three median striae distinct, though 3" stria very short; only finest traces of additional striae vis- ible. Median striae coarsely punctate, on disk impressed, 2™ and 3 behind middle very weak, apical half widely glabrous. Two median intervals on disk slightly raised. 8"
82 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
stria only in apical half present. Recurrent stria deep, elon- gate, oblique, situated close to the lateral margin, at end slightly incurved. Third interval bipunctate, the anterior puncture located at basal quarter and adjacent to 4" stria, the posterior puncture located ın apical third and adjacent to 3" stria. Intervals with a few extremely inconspicuous punctures. Microreticulation absent, surface very glossy.
Lower surface. Prosternum and mesosternum in mid- dle with sparse, very short, erect pilosity. Abdomen not pilose. Metepisternum c. 1.3 times as long as wide at apex. Terminal abdominal sternum in female quadrisetose.
Male genitalia. Unknown.
Female gonocoxites (Fig. 9). Gonocoxite 1 compara- tively large, markedly triangular, without any setae. Gono- coxite 2 fairly elongate, triangular, slightly curved, with fairly acute apex, with two large ventro-lateral ensiform setae and one very elongate subapical nematiform seta that origins from a circular pit.
Variation. Unknown.
Distribution Nepal. Known only from type locality.
Collecting circumstances
Sampled in forested area in river valley but unknown whether by hand collecting or whether at light.
Relationships
In body shape and structure, and in coloration very similar, and probably closely related to 7! quadriplagiata Baehr, 2014.
4.2 Subgenus Eurytachyta n. subgen. Type species: Eurytachyta laticollis, n. sp., here designated.
Etymology
The name of the new subgenus refers to the wide pronotum of the single species.
Diagnosis
Subgenus of Tachyta Kirby, 1837 by virtue of absence of pits on the labrum, finely serrulate tarsal claws, and the elongate, straight recurrent stria that runs close to the lat- eral margin of the elytra.
Special characteristics are the remarkably wide prono- tum with convex lateral margins and excised lateral parts of the base, short and wide, absolutely glabrous elytra, presence of a deeply impressed sutural stria and a very weak 2" stria, total absence of the 8" stria, and insertion of both discal setae at the position of the 3" stria.
The single species Eurytachyta laticollis has been recorded from Sabah, Borneo. For full description of the new subgenus see the description of that species.
Neue Serie 9
In various respects this subgenus differs markedly from the three other subgenera of Tachyta, and likely it could be taken for a separate genus. However, the structure of the recurrent stria, the minutely serrulate tarsal claws, and the narrow and elongate apical palpomere convinced me to include it as a subgenus into the genus Tachyta.
Description of Tachyta (Eurytachyta) laticollis n. sp. (Figs. 10, 18)
Holotype (): “Borneo, Sabah, Kinabalu NP, HQ vic., 1550 m, 22.-25.V.2005, R. Grimm” (CBM).
Etymology The name refers to the unusual width of the pronotum.
Diagnosis A large, wide species, characterized by the very wide, laterally markedly convex pronotum, in the middle strongly produced base and deeply excised lateral parts of the base, and wide, glabrous elytra with a well developed sutural stria and a very weak 2" stria.
Description
Measurements and ratios. Body length: 3.1 mm; width: 1.45mm. Ratios: Width/length of pronotum: 1.62; widest diameter/width of base of pronotum: 1.16; width of prono- tum/width of head: 1.72; length/width of elytra: 1.43.
Colour (Fig. 18). Upper and lower surfaces black, ırı- descent, elytra unspotted, but margins of pronotum and elytra inconspicuously paler. Two basal antennomeres yel- low (rest broken). Palpi and legs yellow.
Head (Fig. 18). Rather wide and short. Eye large, lat- erad well produced, orbit very small, almost perpen- dicular. Labrum anteriorly straight. Clypeus straight, clypeal suture barely indicated. Frons convex, without any impression. Frontal furrows elongate, deep, oblique, slightly surpassing middle of eye. Apical palpomeres elon- gate. Both antennae broken from 3" antennomere. Dor- sal surface impunctate and without microreticulation, very glossy, only labrum with superficial, isodiametric microreticulation.
Pronotum (Fig. 18). Very wide, at base much wider than at apex, strongly narrowed to apex; dorsal surface rather convex. Apex straight, apical angles very widely rounded. Lateral border markedly convex, even near base not concave. Base in middle remarkably produced, lat- eral parts deeply excised, therefore the rectangular basal angles slightly produced posteriad. Lateral sulcus wide and deep, basad slightly widened. Apex completely mar- gined, base laterally margined. Median line very incon- spicuous, not impressed, neither attaining apex nor base.
BAEHR, NEW TACHYINE SPECIES FROM THE ORIENTAL REGION 83
Anterior transverse impression not perceptible, posterior impression deep, smooth, in middle widely interrupted. Basal grooves circular, deep, base near lateral margin with a Short, oblique, not carinate swelling. Anterior lateral seta inserted very close to apex, posterior lateral seta inserted at basal angle. Surface impunctate, without microreticula- tion, very glossy and iridescent.
Elytra (Fig. 18). Moderately elongate, not oviform, lateral margin in middle straight; dorsal surface convex. Humerus evenly rounded. Sutural stria distinct and well impressed, smooth, 2" stria very inconspicous and only visible in basal half, other striae virtually absent. Only the sutural intervals raised. 8" stria completely absent. Recur- rent stria deep, very elongate, oblique, situated close to the lateral margin, at end slightly incurved and ending ina pit. Surface bipunctate, both punctures located at position of 34 stria; the anterior puncture located at basal sixth, the posterior in apical third. Surface impunctate and without microreticulation, very glossy and iridescent.
Lower surface. Prosternum and mesosternum appar- ently glabrous. Abdomen not pilose. Metepisternum slightly <1.5times as long as wide at apex. Terminal abdominal sternum in female quadrisetose.
Male genitalia. Unknown.
Female gonocoxites (Fig. 10). Gonocoxite 1 very large, triangular and globose, without any setae. Gonocoxite 2 fairly elongate, triangular, slightly curved, with acute apex, with two elongate, narrow ventro-lateral ensiform setae, the upper one being longer, one elongate dorso- median ensiform seta, and one elongate subapical nemati- form seta that origins from a circular pit.
Variation. Unknown.
Distribution Sabah, Borneo. Known only from type locality.
Collecting circumstances
Not recorded, but, according to the collector, probably sampled at light.
5 Genus Paratachys Casey, 1918
Paratachys Casey, 1918: 174. — Sctaky & VIGNA TAGLIANTI 2003: 91; Kopecky 2003: 275; LoRENz 2005: 215.
Type species: Paratachys austicinus Casey, 1918, by origi- nal designation.
Diagnosis The genus is characterized by presence of deep mental
pits, not duplicated, curved frontal furrows, smooth tarsal claws, absence of a carina at the basal angle of the prono-
tum, presence of two elytral punctures on disk, interrupted 8" elytral stria, presence of an elongate, at end incurved recurrent stria that is situated rather in the middle of the elytra, and presence of a setiferous puncture right in the curvature of the recurrent stria.
The genus is congruent with the triangularis-group of ANDREWES (1925).
Description of Paratachys leytensis n. sp. (Figs. 11, 19)
Holotype (9): “Philippines, Leyte, visca N Baybay, 1991, sec. forest, 100-200 m, leg. SCHAWALLER & al. / 11.11.1991” (SMNS).
Paratype (9): same locality, but 27.11.1991 (CBM).
Etymology
The name refers to the occurrence of this species on the island of Leyte.
Diagnosis
Characterized by unicolourous, reddish-piceous, markedly iridescent colour, wide, cordiform pronotum, remarkably short and wide elytra, location of the anterior discal puncture at 4" stria, and elongate, straight recur- rent stria with the puncture just inside the short apical curvature. Distinguished from all Oriental and Papuan species with similar structure of the elytra [e. g. P. fascia- tus (Motschulsky, 1851)] by the remarkably short elytra.
Description
Measurements and ratios. Body length: 2.85—2.95 mm; width: 1.30-1.35 mm. Ratios: Width/length of pronotum: 1.60-1.62; widest diameter/width of base of pronotum: 1.26-1.27; width of pronotum/width of head: 1.50; length/ width of elytra: 1.39-1.41.
Colour (Fig. 19). Upper and lower surfaces unicolour- ous reddish-piceous, markedly iridescent. Antenna, palpi, and legs yellow.
Head (Fig. 19). Rather wide. Eye large, but laterad moderately produced, orbit short, oblique-convex, c. '/s of length of eye. Labrum anteriorly straight. Clypeus straight, clypeal suture distinct. Frons convex, without any impression. Frontal furrows elongate, wide, and deep, evenly curved, not duplicated, anteriorly slightly crossing to the clypeus, attaining the posterior margin of the eye. Sulcus medially of the eye deep. Mandible elongate, little curved. Palpi elongate, pilose, apical palpomeres narrow, fairly elongate. Antenna elongate, 7"—10" antennomeres c. 2.25-2.50 times as long as wide. Dorsal surface impunc- tate, with slightly superficial, isodiametric microreticula- tion, moderately glossy but rather iridescent.
84 STUTTGARTER BEITRAGE ZUR NATURKUNDE A
Pronotum (Fig. 19). Wide, at base wider than at apex: dorsal surface gently convex. Apex with fairly deep exci- sion, apical angles projected but widely rounded. Lateral border ın anterior three fourths very convex, in basal fourth oblique and very slightly concave. Base laterally straight, in middle slightly produced, basal angles acute, about 100°. Lateral sulcus rather wide, basad slightly widened. Apex laterally finely margined, base not margined except near basal angles. Median line distinct, slightly impressed, deepened basad,