Bonn zoological Bulletin 64 (2): 59-76 March 2016 Neglected Trichalophus (Coleoptera: Curculionidae): DNA barcode and phylogeography of high-altitude flightless weevils rediscovered in Southwest China Vasily V. Grebennikov Canadian Food Inspection Agency, K.W. Neatby Bldg., 960 Carling Ave., Ottawa, ON K1 A 0Y9, Canada vasily.grebennikov@inspection.gc.ca; v_grebennikov@mail.ru Abstract. Trichalophus LeConte weevils are rediscovered in Southwest China (Yunnan and Sichuan) after last being col- lected in 1915. Populations are all found at high altitudes (3704-4158 m) and are attributed to three species: T. caudic- ulatus (Fairmaire, 1886) (= compressicauda Fairmaire, 1887, syn. n.), T. scylla sp. n. and T. tibetanus (Suvorov, 1915). Type specimens of all four species-group names are illustrated. A DNA barcode library of five Trichalophus species (29 sequences) is presented at doi: dx.doi.org/10.5883/DS-TRICHAL. All examined species of Trichalophus are flightless. Phylogenetic relationships of Southwest China Trichalophus based on Maximum Likelihood and Maximum Parsimony analyses suggest their monophyletic origin and monophyly of each species. Results of the temporal analysis are consis- tent with the basic Quaternary expansion-contraction model of altitudinal range change. The warm period following the Last Glacial Maximum (26,000-19,000 years before present) is linked to the present day high altitude Trichalophus refugium in Southwest China, but not for the lineage diversifications, which are much older (8.08-5. 17 Mya). An illustrated overview of ten extant Alophini genera is provided. Key words. New species, new synonym, Yunnan, Sichuan, COl, Entiminae, Alophini, Father Delavay INTRODUCTION The exclusively Holarctic weevil genus Trichalophus LeConte, 1876 consists of 51 valid species (Yunakov 2013). Adults of Trichalophus are believed to be flight- less and brachypterous (Anderson 1997; Fig. 1). Loss of flight ability is commonly observed in a number of unre- lated edaphic, alpine, high latitude, subterranean, island, or litter- inhabiting insects. Examples from weevils include Macaronesian Laparocerus Schonherr, 1 834 (Machado et al. 2008), VoXynQsmnRhyncogonus Sharp, 1885 (Macha- do 2007), predominantly Oriental Trigonopterus Fauvel, 1862 (Riedel 2011), Middle American Theognete Cham- pion, 1902 (Anderson 2010), an assemblage of likely un- related Old World genera historically placed in Molytini (Grebennikov 2014b) and the Western Palaearctic genera of former “Cryptorhynchinae (Lyal 2014) traditionally linked to Acalles Schoenherr, 1825 (Astrin & Stiiben 2008). The reduced dispersal capacity predisposes such evolutionary lineages to become hostages of their habi- tats and, therefore, subject to more frequent bottleneck ef- fects. These conditions favour allopatric speciation (Grant & Grant 2006; Ikeda et al. 2012; Vogler & Timmermans 2012) seemingly accompanied by an accelerated rate of DNA evolution (Bromham 2008), as compared to their flight-capable relatives (Mitterboeck et al. 2013). Such bi- ological characteristics make flightless weevils a model group for phylogeographical studies on such dynamic ter- Received: 03.02.2015 Accepted: 08.09.2015 rains as oceanic islands (Tanzler et al. 2014; Toussaint et al. 2015) or mountaintops (Grebennikov 2014a), if it were not for their often highly inadequate taxonomy. The lat- ter, if not updated, either lacks names for newly detected evolutionary lineages (= new species), or has too many names for the same clade (= synonyms), or a combina- tion of both. In such situations the historical burden of Lin- naean names impedes, rather than advances evolutionary studies (Riedel et al. 2013a, b). The genus Trichalophus has a trans-Beringian distribu- tion range, with species found on the Pacific sides of both Asia and North America, evoking the Bering land bridge hypothesis (Berman et al. 2011). All but one species are restricted to either Asia or North America; the exception being the Nearctic T hylobinus (LeConte, 1876) recent- ly reported from North Korea (Yunakov 2013). Trans- Beringian distributions are commonly observed in a num- ber of weevil genera: Alaocybites Gilbert, 1956 (Greben- nikov 2010), Thalasselephas Egorov & Korotyaev, 1977, Emphyastes Mannerheim, 1852, Lepyrus Germar, 1817, Lobosoma Zimmermann, 1964, or Lepidophorus Kirby, 1837 (Egorov et al. 1996; Bousquet et al. 2013). Indeed, all but one (Yunakov et al. 2012) Palaearctic records of Trichalophus pertain to Siberia, Mongolia, Central Asia and the northern part of the Pacific Asia including Japan, while the Nearctic species are predominantly found in Alaska, the western USA, and Canada west of Ontario (Anderson 2002; Bousquet et al. 2013). Yunakov (2013) Corresponding editor: D. Ahrens 60 Vasily V. Grebennikov listed 43 Trichalophus species and one non-nominal sub- species {T. vittatoides striola Reitter, 1913) for the Palaearctic region, while Anderson (2002) mentioned eight North American congeners. Since then the Nearctic diver- sity of Trichalophus has lost two species names due to syn- onymy (T. seriatus Mannerheim, 1843 and T. brunneus Van Dyke, 1927) and has gained two others through the recently synonymized genus Acmaegenius LeConte, 1 876 (Bright & Bouchard 2008). As a result, eight Nearc- tic species are currently recognized (T. arcuatus Fall, 1907, T. hylobinusTQConiQ, 1876, T. planirostrisTQCQniQ, 1876, T. seminudus Van Dyke, 1938, T. granicollis Van Dyke, 1927, T. didymus LeConte, 1854, T. simplex LeConte, 1 876, and T. alternatus Say, 1 832), the latter four being recorded from Canada (Bright & Bouchard 2008). Despite the relatively large size of these beetles and their occasional abundance in suitable habitats, biological da- ta on Trichalophus are remarkably scarce. Adult beetles appear highly polyphagous (Anderson 2002) being found on a number of shrubs and herbs (personal observation). North American T didymus was mentioned as an occa- sional pest of strawberries (Fragaria sp.) in Washington State (see references in Bright & Bouchard 2008). Imma- ture stages and larval host plants are adequately known only for a widely distributed Siberian species T leucon Gebler, 1841. Larvae of this species feed externally on the roots of Ribes L. (Grossulariaceae) as well as on a few oth- er shrubs and take two years to complete development (Krivets & Burlak 1986; Krivets 2006). The host plant record is of potential economic significance, since the host genus includes cultivated currants and a number of orna- mental plants. The genus Ribes also includes alternate hosts for the White Pine Blister Rust (Cronartium ribico- la J.C. Fischer, Cronartiaceae), a fungus accidentally in- troduced to North America about 1900 from Europe or Asia, which causes significant damage to American white pines (Pinus spp.; Maloy 2001). Nothing is known about the evolutionary history of Trichalophus and the phylogenetic relationships of this taxon. Flightlessness is not unique for Trichalophus, but it is found in some other genera of Alophini (Bright & Bouchard 2008), including the West Palaearctic Graptus Schoenherr, 1 823 (Davidian & Arzanov 2004). The latter genus has long been known under its synonymous name Alophus Schoenherr, 1 826 and was widely used in origi- nal combinations for Palaearctic Trichalophus prior to Re- itter ’s generic revision (1913). No members of either Trichalophus or any other Alophini were subjected to a phylogenetic analysis so far. The taxonomic recognition of either the genus or the tribe, along with a few diagnos- tic characters used in the keys (i.e. Anderson 2002) are, therefore, the only hints of their possible monophyly. Since the genus Trichalophus was historically linked to Grap- tus, they both might fomi a clade, even if paraphyletic with respect to the North American Plinthodes LeConte, 1 876 (Bright & Bouchard 2008) and perhaps other oligotypic Holarctic genera of Alophini {sensu Alonso-Zarazaga & Lyal 1999, with subsequent modifications of Bright & Bouchard 2008 and Alonso-Zarazaga et al. 2010; see al- so below). DNA data for Trichalophus are exceptionally scarce, with only four public partial COl sequences (>400 bp) of T alternatus presently available from either Bar- code of Life Database (BOLD) or GenBank. LeConte (1876) established the genus for six nominal Nearctic species known to him, four of them having been described earlier as Alophus (A. didymus, A. constrictus LeConte, 1857, A. alternatus, A. seriatus) plus two new- ly described (A. simplex, A. planirostris). Six other cur- rently valid Palaearctic species (Yunakov 2013) described prior to 1 876 were added later; one of them was original- ly described as Hypsonotus Germar {H. boeberi Schoen- herr, 1826) and five others as Alophus (A. albonotatus Motschulsky, 1869, A. humeralis GqHqy, 1834, A. linea- Gebler, 1841, A. quadrigiittatus GqHqx, 1829, A. rud- is Boheman, 1 842). After 1 876, the Palaearctic part of the genus grew quickly in size. By the year 1915 Trichalo- phus included all but six of its 44 currently valid Palaearc- tic species-group taxa (Yunakov 2013). This notable in- crease was mainly due to the efforts of Johannes K.E. Faust and Edmund Reitter who, together with a few oth- ers, introduced 30 currently valid species-group names de- scribed from specimens collected on the Asian frontiers of the rapidly growing Russian Empire (Siberia, Russian Far East, Turkmenistan, Tajikistan, Uzbekistan, Kyrgyzs- tan, Kazakhstan, Mongolia, Xinjiang Uyghur Autonomous Region of China; Pierce 1960; Bajtenov 1974). Oddly enough, two wq^n Alophus species, A. caudiculatus and A. compressicauda, were reported by Fairmaire (1886, 1887, respectively) from geographically distant Yunnan situated on the extreme southwest of China. The latter records, if indeed belonging to Trichalophus, would ex- tend the genus’ range for over 1,000 kilometers south- wards. In 1913 Reitter described 13 species-group taxa in a key to all Palaearctic species known to him. Until now Reitter’s revision has remained the most comprehensive single publication on Trichalophus weevils. For the following hundred years the genus was neglect- ed and only six new species-group names were introduced. Among them are both species recorded from Japan: T rubripes Zherikhin & Nazarov, 1990 and T nutakkanus Kono, 1936; the former also found on the continent, while the latter is endemic to Hokkaido. Two other species were named from the mountains of the former Soviet Central Asia {T. lixomorphus Bajtenov, 1974 and T krauseanus Bajtenov, 1975), while one {T. korotyaevi Zherikhin & Nazarov, 1990) was described from Sakha Republic (=Yakutia). Additionally, Suvorov (1915) established a monotypic genus Pseudalophus for his new species P. ti- betanus described from an unknown number of syntypes collected during Pyotr K. Kozlov’s (1863-1935) Mongol- Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 61 #601 a Tibetan Plateau Sichuan Basin lyanmar Mongolia Russian Far East India approximate southern edge of known continuous Trichalophus range in Asia weevils Is Southwest China: 1 . "Kundur-Tschu rivef : T. f/iwlanus, type locality 2. Mt. Gonnga; T. tibetanus 3. Gang Shan: J. caudicuiatus 4. Mt. Haba; T. oaudicutatus 5- Songpan: T. sp, n. & 7 ?. not localized in "Yunnan*: T. compfassicauda Kazakhstan Kyrgyzstan stub of hind wing Gap of at least 1,000 km Fig. 1. Known geographic distribution of Trichalophus is Southwest China. Inserted image of female specimen #60 1 8 from Kaza- khstan shows hind wing brachyptery in Trichalophus. Tibetan expeditions 1899-1901. The published temporal and geographical data of this species are identical with those of Notaris kozlovi Korotyaev, 1979 (Grebennikov & Kolov, unpublished) and the type locality is in the pres- ent day extreme northwest of Sichuan (Fig. 1). The four- line description of the new genus hinged on two charac- ters distinguishing it from ''Alophus'\ namely (a.) elytra strongly compressed “internally” (i.e. as if a force was ap- plied in the horizontal plane therefore flattening elytra in the vertical plane) and (b.) elytral surface “naked” (= lack- ing vestiture). The former character is known to occur in Palaearctic Trichalophus and was used by Fairmaire (1887) to derive the name compressicauda for a species from nearby Yunnan, while the latter character might per- haps be attributed to abrading. Suvorov’s original descrip- tion was suggestive of a Trichalophus species, and, not surprisingly, Yunakov (2013) synonymized both generic names. This project began on May 19, 2010, when the first two Trichalophus specimens (#0713 & #0714, Figs 3, 5), among those reported below were found under stones in the alpine zone of the Gang Shan Mountain Range in Yun- nan (Fig. 1). The find was most inspiring and seemingly supportive of Fairmaire’s historical claim that the genus was present so far south. During the next two years addi- tional specimens were recovered in the same and three oth- er high altitude localities in Yunnan (Mount Haba) and Sichuan (Mount Gongga and Songpan, the latter seeming- ly supporting two sympatric species). Phenetic similari- ties and subsequent analysis of DNA barcodes suggested that those were indeed species of Trichalophus. The wide gap seemingly separating these Trichalophus of Southwest China from their congeners in the north (Fig. 1) became partly bridged when the former ''Pseudalophus” ti- betanus was transferred to Trichalophus (Yunakov 2013). At that stage it became evident that Trichalophus was in- deed present in Yunnan and Sichuan. The discovery of the high-altitude and the extreme southern representatives of a widely distributed trans-Beringian genus suggests a refu- gial distribution since the last glacial period (Darwin, 1 859: 373-382). The goals of the present paper, therefore, are (1.) to attempt unfolding the evolutionary past of the newly sampled Trichalophus specimens from Southwest China using mainly mtDNA data and (2.) to report these findings in the framework of ranked Linnaean classifica- tion. Additionally, an attempt is made to provide an illus- trated overview of all ten extant genera of Alophini and, therefore, to bring attention to this poorly defined, and tax- onomically disorganized weevil tribe. MATERIAL AND METHODS Museum abbreviations, followed by the curator’s name: CNC Canadian National Collection of Insects, Arach- nids and Nematodes, Ottawa, Canada (P. Bouchard) IZCAS Institute of Zoology, Chinese Academy of Sci- ence, Beijing, PR. China (R. Zhang) MNHN Museum National d’Histoire Naturelle, Paris, France (H. Perrin, A. Mantilleri) Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK 62 Vasily V. Grebennikov Fig. 2. Habitats of Trichalophus spp. in Southwest China. A-C; T. scylla sp. n., Songpan, Sichuan; D-F: T. tibetanus, Songpan, Sichuan; G-I: T. caudiculatus. Gang Shan, Yunnan; J-L: T. caudiculatus, Mt. Haba, Yunnan. Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 63 scyits sp. n. #4437 V #4438 M #4439 M #4440 F #4441 M #5385 F #5386 F #5387 M #5388 F Ffidwifop/tus (jiwlBinius Songpan o-Sb^,7a #2742 F #2743 M #2744 F #2745 M 'Songpien #4418 M #4419 IVt #4436 HHOLOTYPE FnohfffctphfJS Sp. r- #6713 M #0714 F #2767 F #2763 F ICsrg Shafi #4624 M 1 \ #4625 M FxSbaitipfMrs esMfcuffltus Ww1/0 ^ Haba' 0,81 ro, 64 #4620 F #4627 M #5464 M #5405 #5466 F #2960 T aKemaSus Tfictmkipttiji csiiiiiajtBliiS vMble 4 (-slemite 8) venble 5 (=s1emite 7) #5383 F #0714 F #5406F Fig. 3. Maximum Likelihood inference phylogram of Trichalophus weevils from Southwest China using the 658 bp of the mtD- NA barcoding COl gene fragment. The tree is rooted on Graptus circassicus (Entiminae: Alophini; not shown); two extraterrito- rial outgroup Trichalophus (#2960 & #2968) are in grey. Digits at intemodes are ML/MP bootstrap values. Six geographical evo- lutionary groups of Trichalophus are marked as clades A-F in white squares; note that Mount Haba and Songpan each harbours two evolutionary groups. Black long arrows link respective terminal clades with an image of their representative. Specimen num- bers in bold are those of imaged males (M, to the left of the tree) or females (F, in the insert showing ventral view). Black and white short arrows indicate morphological characters and, after a slash, their states (Table 1). Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK 64 Vasily V. Grebennikov MTD Senckenberg Naturhistorische Sammlungen, Dresden, Germany (K.-D. Klass, O. Jager) ZIN Zoological Institute, Russian Academy of Sci- ences, St. Petersburg, Russia (B.A. Korotyaev) The length of the body was measured in dorsal aspect from the elytral apex to the anterior edge of the pronotum. Dis- tribution map (Fig. 1) is generated using the on-line Sim- pleMappr tool (Shorthouse 2010). The chronostrati- graphic timing follows Cohen et al. (2013). Nomenclature of male genitalia follows that of Wanat (2007). The term “Southwest China” is delimited to two Chinese provinces, Yunnan and Sichuan. The term “base pair” is abbreviat- ed as bp when referring to sequence length; abbreviations “syn. n” and “sp. n.” denote new synonym and new species, respectively. Specimen sampling, handling and gathering DNA data Except for two specimens #4418 and #4419 sifted from Rhododendron L. leaf litter, all newly collected Trichalo- phus were handpicked from under stones (Figs 2A-L) in the alpine zone (Fig. 2J), or on a glade in the upper for- est zone (Fig. 2D) in Southwest China. In total 38 adult Trichalophus beetles were collected in the following four localities (Fig. 1 ; in brackets are the total number of spec- imens followed after a slash by the number of those suc- cessfully sequenced for DNA barcode >400 bp): the Cang Shan Mountain Range (4/4), Mount Haba (12/7), Mount Gongga (4/4), and the vicinity of Songpan township (1 8/12). A leg was removed from a specimen for DNA ex- traction. All specimens used for DNA barcoding have at least one unique identifier label with the code CNC- COLVGOOOOXXXX; this format is shortened to the last four digits #XXXX when a specimen is referred to (Figs 3,4). Specimen images, geographical data, primers, orig- inal electropherograms and other relevant data pertaining to all 35 matrix-forming sequences can be seen online in the publicly accessible dataset ''Trichalophus 35 [DS- TRICHAL]” on the Barcode of Life Database portal (doi: dx.doi.org/10.5883/DS-TRICHAL). Genitalia of six males each representing a terminal cluster (=evolutionary group) as detected in the phylogenetic analyses (see below) were dissected, imaged (Fig. 3) and stored in microvials with glycerol pinned with the specimens. DNA analyses and matrix constrnction Three separate DNA analyses were performed. The Max- imnm Likelihood (ML) and the Maximnm Parsimony (MP) analyses attempted to place the diversity of Table 1. Discrete morphological characters for diagnostics of Trichalophus weevils in Southwest China (Fig. 3) 1. Elytral shoulders and elytral sides in basal 2/3, dorsal view; shoulders rounded, sides evenly widening posterad (0); shoulders angular, sides subparallel (1). 2. Elytral dorsal and posterior profile (=dechvity), lateral view; evenly and gently rounded throughout (0); flattened dorsally and abmptly curved (1). 3. Female, ventrite 5, bumps and depressions on surface, ventral view; absent (0); present (1). 4. Female, ventrite 4, two sharp points at posterior edge, ventral view; absent (0); present (1). 5. Female, posterior projections of elytral apices, ventral view: absent (0); present (1). 6. Male genitalia, long, thin, and curved apical lamella of aedeagus (lateral view); absent (0); present (1). 7. Male genitalia, aedeagus, dorsal view: symmetrical (0); asymmetrical (1). 8. Male genitalia, aedeagus, notch in lateral outline, dorsal view; absent (0); present (1). 9. Male genitalia, sternal apodeme 9, dorsal view; thin (0); thick (1). clade species Locality 1 2 3 4 5 6 7 8 9 A tibetanus Songpan 0 0 0 1 0 1 0 1 0 B tibetanus Mt. Gongga 0 0 0 1 0 1 0 1 0 C scylla sp. n. Songpan 0 0 7 7 7 0 1 0 1 D caudiculatus Cang Shan 1 1 1 0 1 0 0 0 1 E caudiculatus Mt. Haba 0 1 7 7 7 0 0 0 1 F caudiculatus Mt. Haba 0 1 1 0 1 0 0 0 1 Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 65 Trichalophus from Southwest China into phylogenetic and geographical perspective. Both ML and MP used the same matrix of 35 DNA barcodes with a minimum and maxi- mum length of 400 bp and 658 bp, respectively. The in- group consisted of 27 Trichalophus specimens from Southwest China, while the outgroup included two extra- territorial Trichalophus specimens representing T rubripes from the Russian Far East and T alternatus from Cana- da (Fig. 3). The rest of the outgroup was formed by six specimens of Graptus circassicus Solari, 1945 from Geor- gia: Abkhazia. Both ML and MP analyses were imple- mented using MEGA 6 (Tamura et al. 2013), including (a.) topology building, (b.) statistical support test by using 1000 repetitions of bootstrapping and (c.) search for the best substitution model for ML analysis (T92+G). The root was consistently placed between Graptus and Trichalo- phus. The GenBank accessions for these 35 sequences are KM538655-86, KJ445708, KJ445709, KJ445712; all of them are new, except for the three latter Graptus se- quences. The third DNA analysis was performed to date the in- group branching events and to re-test the phytogenies sug- gested by the ML and MP analysis. The original matrix of 35 sequences was reduced to include only 23 full-length DNA barcodes (658 nt; except for the sequence of T al- ternatus with 609 bp). The second analysis was performed in BEAST vl.8.0 (Drummond et al. 2012) utilizing the Bayesian inference (BI) approach with no a priori group- ing, all default priors and options, GRT+G+I nucleotide substitution model (instead of the T92+G+I not offered in the software; the latter model was detected in a separate model- searching analysis in MEGA 6 as having the best fit), strict linear molecular clock and nucleotide substitu- tion rate of 0.018 (Papadopoulou et al. 2010). Tracer 1.6 (Rambaut et al., 2014) was used to graphically determine stationarity and to check convergence of runs. The “burn in” option was implemented eliminating the first 2500 of the 10000 obtained trees. The resulting topologies from each of three analyses (ML, MP, BI) were visualized in FigTree vl.4 (Rambaut et al. 2014). Contribution from morphology Morphological data are not expected to contribute deci- sively in DNA-dominated phylogenetic analysis, partic- ularly in shallow branches of the tree of life (Ward 2011) conventionally called “species” in ranked classification (Hey 2001). Consequently, no effort was made to merge the DNA matrix with a few morphological characters scored for the ingroup (Table 1). Instead, DNA-determined Trichalophus clusters (= evolutionary groups by Hey 2001 or cladesA-F on Fig. 1) were a posteriori scrutinized in search of diagnostic morphological characters (Maddison 2014), not necessarily synapomorphic (Ward 2011). The easily observable dorsal color pattern, being either too variable or subject to abrasion, was judged as unreliable for diagnostic purposes in Trichalophus. An additional ef- fort was made to explore structures of male genitalia by dissecting a single male per each of six clades detected on the phylogenetic tree (Fig. 3). Integrating molecular phylogenetic results into taxonomy In biodiversity studies taxa are normally named first and then their phylogeny and boundaries are analysed, if ever. This approach, although logically awkward, has strong his- torical roots from the times when (a.) phylogenetic theo- ry was not practiced by taxonomists, and (b.) researchers lacked adequate data to perform sufficiently detailed analyses when naming their new species. Advent of Hen- nigian principles coupled with availability of DNA se- quences challenges this classical and logically deficient approach (Ward 201 1). A modern student of biodiversity is expected to (A.) delimit evolutionary groups through a formal analysis, then (B.) make a balanced, responsible and subjective judgement using all available evidence sources as to which clades need names (Hey 2001) and then (C.) conservatively apply formal names, either pre- existing or newly proposed. In other words whenever pos- sible, taxa naming should not be done before but after the analysis and discussion, not to abuse logic by putting the cart before the horse. Through most of the present paper all six tree-delimited clades representing candidate species (Fig. 3) are referred to by using informal non-taxonomic names (clades A-F, italicized). Therefore, the taxonomic part of this paper using three valid species-group names (two previously used and one new) and synonymizing one name follows the Results section and most of the Discus- sion. RESULTS The Maximum Likelihood (ML) analysis produced the best tree (Fig. 3) with the highest log likelihood of -3265.92. All Trichalophus specimens from Southwest China formed a weakly supported clade with highly re- solved internal structure consisting of six clades A-F. The Maximum Parsimony (MP) analysis resulted in seven best trees (length: 558, consistency index: 0.64, retention in- dex: 0.90; topologies are not shown) also recovering the same six clades A-F (as in Fig. 3). In both analyses the ingroup was recovered as a clade. The only backbone topological difference of the MP strict consensus tree (as compared to the ML topology. Fig. 3) was that the clade C was recovered as the sister to the rest of Trichalophus from Southwest China. Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK Vasily V. Grebennikov Pliocene Pleistocene T rubhpes 10 6 0 Fig. 4. Ultrametric tree dating evolutionary events of Trichalophus beetles in Southwest China. Digits at nodes and on the scale below are million years before present. Node bars represent 95% confidence intervals of the age estimate. Temporal analysis in BEAST (Fig. 4) recovered all Trichalophus specimens from Southwest China in a clade with the same internal backbone topology as in the ML analysis (Fig. 3, although some specimens and the entire clade E were not represented in the BEAST analysis due to inadequate sequence length). The inferred timing of the origin of the Southwest China Trichalophus is 1 1 .65 MY, while the clade’s diversification leading to the five clades A-D and clade F range between 8.08 MY and 5.17 MY (Fig. 4). DISCUSSION mtDNA phylogeny and phylogeography of Trichalophus in Sonthwest China Recovery of a monophyletic Trichalophus radiation in Southwest China (Figs 3, 4) should be treated with cau- tion, since limitations in the number of the in- and out- group representatives did not permit a rigorous test. Lit- tle other evidence is available to challenge this hypothe- sis. The entire clade A-F has a compact range allopatric to that of the rest of the genus (Fig. 1), although the dis- junct distribution might be plausibly attributed to the lack of adequate sampling effort to bridge the gap. Morpho- logically Trichalophus beetles from Southwest China are seemingly large-bodied compared to other two species (Fig. 3), although adding more Trichalophus in the analy- sis might challenge this pattern. Nearly all of the ingroup morphological characters (Table 1) cannot be adequately matched with those for the rest of the genus due to the lack of comparative data. Summing up, compact distribution (Fig. 1) of monophyletic Trichalophus of Southwest Chi- na (Figs 3, 4) is a weakly supported hypothesis to be even- tually retested. Internal structure of six ingroup clades (Fig. 3) is mainly consistent with limited geographical (Fig. 1) and morphological (Table 1) data and is further discussed when Linnaean species are delimited (see below). Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 67 Syntypa of Syn^M of coimressvcaLic Vi totl . IbiJiia Firli :»u. It. C(f*rtliiiT ^ l«2 rOHiln (X&pytj 1 Vfi. $tt^O-rQV duL Fig. 5. Type specimens and original labels of three historical Trichalophus species names from Southwest China {caudiculatus, compressicauda, tibetanus currently assigned to two valid species T. caudiculatus and T. tibetanus), together with three T. caudic- ulatus females (#0714, #2767, #27678) sequenced for DNA barcode (Fig. 1). Black and white arrows indicate morphological char- acters and their states (Table 1, separated by a dash). Images of caudiculatus and compressicauda syntypes and their labels: An- toine Mantilleri, © MNHN. Unlike at least some of its more northern congeners, Trichalophus in Southwest China inhabit high altitudes (3704-4158 m). Such a characteristic of the southern-most representatives of a temperate northern hemisphere clade of low-dispersing organisms is consistent with the basic Quaternary expansion-contraction model of latitudinal range change (Qiu et al. 2011). The latter stipulates ex- tensive latitudinal range shifts in the form of southward movement during glacials followed by rapid expansions northwards during interglacials (Qiu et al. 2011). If so, dis- tribution of Trichalophus in Southwest China (Fig. 1) is a direct result of the last warming following the Last Gla- cial Maximum (26,000-19,000 ybp). More specifically, the observed data are consistent with at least three sub- hypotheses (numbering after Qiu et al. 2011): (iii) long- term isolation and survival in multiple localized refugia (cladesA-D and clade E+F in Fig. 3), (ii) population iso- lation and endemism due to river course dynamics {clade D versus clade E+F) and (iv) glacial in situ survival of some hardy alpine species on the Tibetan plateau itself (population of T tibetanus represented by the type spec- imens, Fig. 5). Like the hypothesis on monophyly of Trichalophus in Southwest China, all phylogeographical inferences are highly preliminary due to material and da- ta limitations. With no suitable fossils to calibrate a Trichalophus mo- lecular clock, the temporal aspect of Trichalophus evolu- tion in Southwest China (Fig. 4) relies on the a priori mtD- Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK Vasily V. Grebennikov NA substitution rate of distantly related Tenebrionidae (Pa- padopoulou et al. 2010). The obtained dates of the Trichalophus cladogenesis (Fig. 4) are comparable to those of the sympatric and similarly high-altitude and flightless weevil genus Niphadomimus Zherikhin, 1987 (Greben- nikov 2014a). Both agree that the lineage divergence took place well before the onset of the Pleistocene climate fluc- tuations. Both time estimations were based, however, on identical methods and substitution rates, which might have biased them both. Exact substitution rates may significant- ly vary depending on population size, founder effects, and a number of other less well-understood factors and, there- fore, markedly differ from the assumed 0.018 substitutions per site per million years. In other words application of the user-friendly BEAST software and the obtained clear- cut dates (Fig. 4) should be treated carefully, since we are far from understanding the molecular clock (Eanfear et al. 2010), even for such relatively simplified and recent sce- nario as that of Trichalophus weevils in Southwest Chi- na. Delimitation of Linnaean species for Trichalophus in Southwest China The practical task of assigning Einnaean species names to the newly discovered Trichalophus from Southwest China, even with the help of a tree (Fig. 3), is far from being trivial. It involves at least one theoretical and two practical difficulties. First, imposing ranked classification on the continuum of the tree of life cannot be objectivised (Hey 2001 ; Ward 2011) and, therefore, involves arbitrary decisions (Sites & Marshall 2004). Second, despite sam- pling and analytical efforts, relatively little data on Trichalophus are available. For example, the relatively well-resolved DNA tree (Fig. 3) is that of a very short frag- ment of a fast-evolving mitochondrial maternally inher- ited gene and, therefore, only a proxy to organismal evo- lutionary history. Third difficulty is the existence of three available historical names, which have nomenclatorial pri- ority and have to be interpreted and used, if considered as valid. These three issues have to be considered before the freshly sampled Trichalophus in Southwest China might be incorporated into the existing taxonomic scheme (Yunakov 2013). The first difficulty, concerning the issue of what a Trichalophus “species” is, will be resolved in this passage, while both practical issues are considered fur- ther below. A “species” as a taxonomic category is purely and sole- ly a label routinely and often inconsistently assigned to the shallow branches of the tree of life ever since Einnaeus. As such, “species” is no more real than other taxonomic categories like “genus”, “family” or “phylum” (Hey 2001). Eike every other taxonomic category, a “species” is a mere convenience required by the human mind to categorize and count biological diversity. Acceptance of this basic philo- sophical and methodological principle denies “species” re- ality in the same sense at it denies reality “genera” or oth- er “higher” taxonomic categories. In practical terms a species is nothing more than a morphologically (or oth- erwise) diagnosable group of organisms preferably form- ing a clade and, most importantly, considered practically worthy of being called a species (Hey 2001). If to follow such an approach, decision on species boundaries almost fully rests with the revising author, which, in turn, results in the splitters versus lumpers issues, particularly in re- gard to a clade of allopatric populations. The authors nor- mally feel free to either split them into as many species as possible, or lump them into a single one. The first ap- proach is an example of unnecessary taxonomic inflation (Isaac et al. 2004) adding nothing but unnecessary names. The alternative lumping approach using a single species name and a geographic reference would label every al- lopatric evolutionary group equally well, while avoiding unnecessary additions to the already heavy nomenclato- rial burden. Moreover, a scramble to call a “species” an allopatric evolutionary group, even if accompanied by cor- relating morphological and other differences, can be mis- leading when linked to the phenomenon of Sisyphean evo- lution (McKay & Zink 2014). Summing up, introduction of new species names should be done only when all al- ternative options have been shown as inadequate. Such a careful and reserved approach would not have created the multitude of meaningless and cryptic taxa (Riedel et al. 2013a, b; Vences et al. 2013). Two among six terminal clades representing Trichalo- phus evolutionary groups (Fig. 3) are allopatric to all oth- er ingroup clades {clade B and clade D from Mt. Gong- ga and Gang Shan, respectively). They should, therefore, be first assessed if each of them can be merged together with its strongly supported sister-group into a more inclu- sive clade to merit a species name. Indeed, clade B is strongly linked to the allopatric clade A (Fig. 3), so the clade A+B might itself be considered a candidate species. Clade C is recovered in the ME analysis as a sister to clade A+B (Fig. 3), but in MP analysis the sister group of clade A+B was the clade D+E+F. Remarkably, specimens of both clade A and clade C occur in sympatry in Songpan (or at least in parapatry; their two geographically closest samples were taken a few hundred meters apart). More- over, while males and females of both clade A and clade B share all eight morphological character states (Table 1), their males differ in three genitalia characters from those of clade C. Such considerations strongly suggest that clades A+B and clade C (Fig. 3) should be treated as two separate species, respectively, and the former one as con- sisting of at least two geographically and morphological- ly unique evolutionary groups {clade A and clade B, Fig. 3). It is possible that in the future each clade A and clade B might be considered as separate species, but not until Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 69 the current nomenclatorial lumping arrangement is refut- ed as impractical. The clade D+E’+F emerges as the third and the last one to be designated as a Linnaean species among those rep- resented in the ingroup (Fig. 3). Remarkably, clade E and clade F from Mount Haba, although genetically distinct, are formed by morphologically and geographically indis- tinguishable specimens (Fig. 3). This notable genetic di- morphism accompanied by full sympatry and morpholog- ical similarity is perhaps linked to incomplete lineage sort- ing (Funk & Omland 2003). The clade E+F is strongly supported as a sister to the geographically closest clade D and both share all but one morphological character (Table 1). Like the clade A+B, the clade D+E+F 3) might also later require two Linnaean species, but not be- fore the present conservative nomenclatorial decision is shown as inadequate. Matching historical names with the tree-delimited Trichalophus Linnaean species The most significant practical constraint is how to link three clades delimited for designation as Linnaean species (clade A+B, clade C and clade D+E+F, Fig. 3) with three available historical names (T. caudiculatus, T. compres- sicauda and T. tibetanus). The type specimens of the lat- ter are well preserved and available for study (Fig. 5). Matching the type specimens of three historical names with three evolutionary groups in Fig. 1 can be done us- ing three sources of evidence: (a.) similarity in body shape and in male genitalia, (b.) geographical proximity and (c.) biological characteristics expressed through the altitude of the type locality. No attempt was made to extract DNA from the type specimens, mainly because they were judged too old to warrant an attempt. Of the three historical names, only the type specimen of T. tibetanus has information available from all three sources. Its type locality can be traced precisely (Figs 1, 5), while habitus and shape of male genitalia of a syntype (Fig. 5) match most closely those of the clade A+B (Fig. 3). The name T. tibetanus is, therefore, used to designate the clade A+B (Fig. 1). Such matching is far less straight- forward with the both Fairmaire’s names. The most significant uncertainty with both names cau- diculatus and compressicauda is that their type localities are imprecise, originally given as “Yunnan”. The years when the types were collected are also unknown. Only the younger of these two names, compressicauda, has the col- lector’s name stated: Pere Jean Marie Delavay (=Father Delavay). It seems, however, plausible that Father Delavay also collected the type series of caudiculatus. Even though Fairmaire cited Armand David (= Father David) as the type specimen source for species described together with caudiculatus in the 1886 paper. Father Delavay was likely the original collector of at least some of them, since Fairmaire in the same paper also named Ci- cindela delavayi Fairmaire, 1886 in his honour. Indeed, the years before both species were described (1 886, 1 887) correspond with Delavay’s second stay in China (1882-1891) (Anonymous 2014). This trip took place af- ter Father Delavay’s meeting with Father David in 1881, the latter convincing the former to collect specimens for the Museum national d’histoire naturelle (Anonymous 2014). During his second stay in China, Delavay was mainly based in a Yunnan village somewhere between Lake Erhai and Eijiang township given as “Dapingzi” (Anonymous 2014). Delavay had two favourite climbing spots nearby: “Mount Heishanmen” (or “Ma’an Shan”, west of “Dapingzi”, see Handel-Mazzetti 2014; not def- initely located, but distinct from the Cang Shan Mountain Range) and the Cang Shan Mountain Range along the western shore of lake Erhai (Eancaster 1993, Anonymous 2014). Besides these two high mountain localities. Father Delavay is definitely loiown to have collected in the alpine zone around Deqin in northwestern Yunnan, from where numerous alpine Carabus were sampled (T. Deuve, per- sonal communication). It is highly probable that Father Delavay visited many other alpine localities in Yunnan, however those three were apparently most frequently vis- ited and/or sampled for high altitude beetles. Both caudiculatus and compressicauda were described from an unknown number of syntypes. Curatorial search- es in MNHN in 2014 revealed a single syntype for each for these names (Antoine Mantilleri, personal communi- cation). Their syntype status is corroborated by the fact that both specimens bear original identification labels and, moreover, fully agree with relatively detailed descriptions of both nominal species (Antoine Mantilleri, personal communication). Both syntypes, although not dissected, appear to be females by possessing posterior projections on elytral apices (character 5/1, Fig. 5) and posterior pro- jections of ventrites 4 (character 4/1, Fig. 5). These two syntype characters match with those of the female spec- imens from clade D (Fig. 3); the other being angular ely- tral shoulders and subparallel elytra in their basal 2/3 (Table 1). Specimens from clade D inhabit the Cang Shan Mountain Range, which was visited on many occasions by Father Delavay, including altitudes above 4,000m (Ean- caster 1993). Both Fairmaire’s names, therefore, represent the best fit for the Einnaean species represented on Fig. 3 by the clade D+E+F, which, in turn, suggests their syn- onymy proposed below. Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK 70 Vasily V. Grebennikov Taxonomic overview of Trichalophus in Sonthwest China Trichalophus LeConte, 1876 Type species: Alophus didymus LeConte, 1854, fixed by subsequent designation (Bright & Bouchard 2008: 57). Trichalophus caudiculatus (Fairmaire, 1886) (Figs 2, 3, 5) caudiculatus YdLixmmxQ 1886: 353 {Alophus) Type locality. Yunnan. Type specimens. Syntype (MNHN, Fig. 5), likely female, examined from images in Fig. 5, labels in Fig. 5. De- scribed from unknown number of syntypes. = compressicauda Fairmaire, 1887: 129 {Alophus) syn. n. Type locality. Yunnan. Type specimens. Syntype (MNHN, Fig. 5), likely female, examined from images in Fig. 5, labels in Fig. 5. De- scribed from unknown number of syntypes. Diagnosis. This species is recognized by presence of api- cal elytral projections in females (character 5/1, Fig. 3). Intraspecific variation. GenBank accessions: KM538662, KM538665, KM538667-68, KM538670-71, KM538676-77, KM538681-82, KM538684. Length: 12.3-14.5 mm (Gang Shan), 11.1-11.8 mm (Mount Ha- ba, clade E) and 11.4-14.5 mm (Mount Haba, clade F). Specimens from both sampled localities (Fig. 1) slightly differ in dorsal coloration, shape of elytral shoulders and arrangement of posterior projection on ventrite 4 of fe- males (Fig. 3). Aedeagus of a single male known from the Gang Shan Mountain Range is less curved and more elon- gate, as compared to those of two males dissected from Mount Haba (Fig. 3), while female posterior projections on elytral apices of the Gang Shan specimens are notice- ably longer than those from Mount Haba (Fig. 3). Additional material examined. 16 exx in total: 2 exx #0713-0714 (GNG) “P.R. GHINA, Yunnan, E slope Gang- shan at Dali, N25°39’54.7” E100°06’04.5”, 19.V.2010, 3815m, turn rock, V. Grebennikov”; 2 exx #2767-2768 (GNG) “RR. GHINA, Yunnan, Gang Shan at Dali, N25°39’51” E100°06’05”, 04.vii.2011, 3815m, under stone, V. Grebennikov”; 12 exx #4623^628, #5403-5406, #6207-6208 (GNG) “GHINA, Yunnan, Haba Shan, N27°20’51” E100°05’33”, 27.vi.2012, 4158m, under rock, V. Grebennikov”. Distribution. This species is known from the Gang Shan Mountain range and nearby Mount Haba, both in Yunnan (Fig. 1). Elevation: 3815^158 m. Trichalophus scylla sp. n. (Figs 2, 3) Diagnosis. Specimens of this species are unique among known congeners in Southwest Ghina by two characters: they are the smallest and possess asymmetrical aedeagus in dorsal view (character 7/1, Fig. 3). Description. Holotype, male (Fig. 3). GenBank accession: KM538655. Eength: 9.4 mm. Gombination of other mor- phological characters as in Table 1 . Intraspecific variation. GenBank accessions: KM538656, KM538679. Eength: 9.0-9.4 mm. Material examined. Holotype (Fig. 3) male (IZGAS): #4436: “GHINA, Sichuan, 23km E Songpan, N32°38’07” E103°49’10”, 24.V.2012, 3704m, under rock, V. Greben- nikov”. Paratypes (GNG): 2 males #4418 and #4419 “GHI- NA, Sichuan, 23km E Songpan, N32°37’38” E103°50’03”, 26.V.2012, 3791m, sifting 09, V. Greben- nikov”. Distribution. This species is known only from the type locality some 20 km E of Songpan, Sichuan (Fig. 1), where it is found sympatrically with T. tibetanus. Eleva- tion: 3704-3791 m. Etymology. The species epithet is the Eatinized Greek mythical name of Scylla, one of the Nereids, transformed by Girce into a six-headed monster and who, together with its counterpart Gharybdis, threatened Odysseus’ crew on their return voyage from Troy to Ithaca; noun in apposi- tion. Trichalophus tibetanus (Suvorov, 1915) (Figs 2, 3, 5) tibetanus Suvorov 1915: 338 {Pseudalophus) Type locality. Ghina, basin of the Blue river (=the Yangtze), the Kundur-Tschu river, 13200’. Type specimens. Syntype (ZIN, currently on loan in MTD, Fig. 5), male, dissected by Rudiger Krause, labels as in Fig. 5. Described from unlmown number of syntypes. Twenty other similar specimens collected together with the imaged syntype are also likely part of the type series; of them four specimens each have a golden circle as in Fig. 5. Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 71 Diagnosis. This species is best distinguished by the pres- ence of elongate and curved apical labella of aedeagus (character 6/1, Fig. 3) and relatively thick apodeme of male sternite 9 (character 9/0, Fig. 3). Intraspecific variation. GenBank accessions: KM538657, KM538658, KM538659, KM538661, KM538663-64, KM538672-75, KM538680, KM538685-86. Length: 11.5-13.3 mm (Songpan) and 10.3-12.1 mm (Mount Gongga). Each elytron with sin- gle indistinct apical spot (Fig. 5), two indistinct spots, or two distinct spots (Fig. 3). Specimens from Mount Gong- ga have a long white longitudinal stripe laterally on each elytron (Fig. 3). Additional material examined. 19 exx in total: 4 exx #2742-2745 (CNC) “RR. CHINA, Sichuan, NE slope Gongga Shan, N25°53’53” E102°0r49”, 8.vi.2011, 4085m, under stone, V. Grebennikov”; 15 exx #4437-4441, #5385-5388 and six not numbered speci- mens in ethanol (CNC) “CHINA, Sichuan, 23km E Song- pan, N32°38’07” E103°49’10”, 24.V.2012, 3704m, under rock, V. Grebennikov”. Distribution. Besides the type locality in the extreme north-western Sichuan, this species is also known from Mount Gongga and from vicinities of Songpan, both in Sichuan (Fig. 1); in the latter locality this species is found sympatrically with T. scylla sp. n. Elevation: 3704^085 m. Generic overview of the tribe Alophini The proposed amalgamation of Alophini (as delimited be- low) with the tribe Tropiphorini {sensu Alonso-Zarazaga & Eyal 1999) by Zherikhin & Egorov (1991), for which a synonymous name Eeptopiini was used by Marvaldi et al. (2014), is not followed here. Neither taxonomic arrangement was phylogenetically tested, therefore none is better than the other. Additionally, dissolving the com- pact and predominantly Holarctic Alophini in the much larger cosmopolitan Tropiphorini would discourage any practical effort to shed light on the genera involved, as at- tempted below. The tribe Alophini was first proposed by EeConte (1876: 115) to incorporate the Palaearctic species grouped then in Alophus Schoenherr and species belonging to sixNearc- tic genera, five of them newly established: Triglyphus EeConte, Plinthodes EeConte, Acmaegenius EeConte, Trichalophus EeConte, Lophalophus EeConte and Lepi- dophorus Kirby, 1837. By the end of the millennium the tribe consisted of 15 valid genera, including three de- scribed from the Oligocene of the USA (Centron Scud- der, 1893, Ger alophus Scudder, 1893 and Limalophus Schudder, 1893, see Alonso-Zarazaga & Eyal 1999). Since then Bright & Bouchard (2008) synonymised the genus Acmaegenius under Trichalophus and reviewed the Alophini of Canada and Alaska. Alonso-Zarazaga et al. (2010) demonstrated that the sole known specimen of the monotypic genus Ctenolobus Desbrochers des Eoges, 1 892 from Morocco is conspecific with the type species of the otherwise strictly South American genus Stran- galiodes Schoenherr, 1842 (Tropiphorini). As a result, Strangaliodes was transferred to the otherwise strictly Palaearctic Alophini and was keyed out against three oth- er Mediterranean genera: Graptus, Rhytideres and Seidl- itzia (Alonso-Zarazaga et al. 2010). Finally, Yunakov (2013) synonymized Pseudalophus under Trichalophus. At present the following 10 extant genera constitute the tribe Alophini: Graptus Schoenherr, 1823 (Figs 6A, B) with 37 species-group taxa is distributed in Western Palaearctic (Yunakov 2013). Davidian & Arzanov (2004) revised and keyed 10 Graptus species from Russia and adjacent lands, including two newly described ones, and mentioned that many poorly known nominal species have been reported from the West Palaearctic. Lepidophorus Kirby, 1837 (Figs 6C, D) consists of 11 brachypterous species found in western North America (Anderson 1997, 2012; Bright & Bouchard 2008), two of which, Z. inquinatusMmnQxhQim, 1852 andZ. lineaticol- lis Kirby, 1837, are also found on the Asian side of the Bering Strait (Yunakov 2013). Another extant North American species, Z. thulius Kissinger, 1974, is known from numerous subfossil records on both side of the Bering Strait and, therefore, extant populations might per- haps be discovered in the northern Pacific Asia (as Vitat- itiis Kissinger, 1974 in Egorov et al. 1996 and in Ander- son 1997). Anderson (2002) mentions that the genus can- not be reliably distinguished from Dirotognathus Horn, 1876 (Tropiphorini Alonso-Zarazaga & Eyal 1999), giving support to a notion to synonymize both tribes (Zherikhin & Egorov 1991; Marvaldi et al. 2014). Plinthodes LeConte, 1876 (Figs 6E, F) consists of two North American species, P.foveirostris Chittenden, 1925 from Ohio, North Carolina, Tennessee and Virginia and P taeniatus EeConte, 1857 from British Columbia, Wash- ington and Oregon (Anderson 2002). Bright & Bouchard (2008) questioned the distinctness of this genus from Trichalophus. Pseudobarynotus Desbrochers des Loges, 1891 (Fig. 8) contains a single mysterious species P. laticeps (Des- brochers des Eoges, 1875) known only from the type se- ries and described from “Pyrenes”. The type series could have been mislabelled, while Kazakhstan was suggested as its true origin (Alonso-Zarazaga et Eyal 1999). The lat- er possibility is not unlikely, since the depicted syntype (Fig. 8) resembles a species of Trichalophus and might perhaps be later demonstrated as such. Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK 72 Vasily V. Grebennikov Fig. 6. Type species of the Alophini genera. A-B; Graptus triguttatus (Fabricius, 1775), Austria, Wien, date and collector un- known, CNC; C-D: Lepidophorus lineaticollis Kirby, 1 837, USA, Alaska, Wasilla, 1 .viii. 1988, J.Pilny, CNC; E-F ; Plinthodes tae- niatus LeConte, 1857, Canada, British Columbia, Victoria, 6.vii.l962, B.Carr, CNC; G-H: Rhytideres plicatus (Oliver, 1790), no collecting data, CNC. Scale; 2 mm. Rhytideres Schoenherr, 1823 (Figs 6G, H) includes three species widely distributed around the Mediterranean Sea (Yunakov 2013). Seidlitzia Desbrochers des Loges, 1891 (Figs 7A, B) consists of two species and one non-nominal subspecies from Spain and Morocco (Yunakov 2013). Strangaliodes Schoenherr, 1842 (habitus image in Alonso-Zarazaga et al. 2010, figs lA, B) includes nine species from the South American Cordillera, all of them found in Chile and a few in neighbouring countries (Wib- mer & O’Brien 1986); one of them also questionably recorded from Morocco (Alonso-Zarazaga et al. 2010). This is the only non-Palaearctic member of Alophini. Trichalophus LeConte, 1876 (Figs 7C, D) consists of 51 species and one non-nominal subspecies distributed on both sides of the Bering Strait; for more details see the current paper. Triglyphulus Cockerell, 1906 (Figs 7E, F) consists of two species, T. ater LeConte, 1 876 and T. nevadensis Van Dyke, 1938 from the western part of the USA (Anderson 2002 ). Xeralophus Korotyaev, 1992 (Figs 7G, H) was estab- lished to accommodate Alophus cretaceus Reitter, 1 894, described from present day Ulan Bator, Mongolia. Besides Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 73 Fig. 7. Type species of the Alophini genera. A-B: Seidlitzia maroccana (Fairmaire, 1 868), Morocco, date and collector unknown, MNFIN, image: Antoine Mantilleri, © MNHN, original image showing right lateral view was digitally flipped horizontally to ap- pear as left; C-D; Trichalophus didymus (LeConte, 1854), Canada, British Columbia, Kitsumkalum Lake, 16. vi. 1960, B.S.Hem- ing, CNC; E-F; Triglyphulus ater (LeConte, 1 876), USA, Idaho, Bear, 26.vii. 1977, B.Carr, CNC; G-H: Xeralophus cretaceus (Re- itter, 1894), Russia, Tyva, Kyzyl, 6.V.1974, B.A.Korotyaev, ZIN, image: Audrey Frolov. Scale: 2 mm. the type series, four more specimens were later reported, all found dead in sandy steppe of the neighbouring Tyva Republic of Russia (Korotyaev 1992). Korotyaev (1992) hypothesised that this xerophilic taxon is phylogenetical- ly nested within the predominantly mesophilic '"Alophus” (= Graptus). This hypothesis, if corroborated, would ren- der the name Xeralophus a junior subjective synonym of Graptus. CONCLUDING REMARKS It seems worthy of reiterating some important and perhaps not too obvious generalities emerging from this study. First, if the genus Trichalophus does exist in the phylo- genetic sense, then its presence has been reconfirmed in Southwest China for the first time since Suvorov (1915). Second, the last glacial retreat is likely responsible for the Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK 74 Vasily V. Grebennikov Fig. 8. Pseudobarynotus laticeps (Desbrochers, 1874), syntype, MNHN, image: Antoine Mantilleri, © MNHN. present day high altitude presence of Trichalophus in the high mountains of Yunnan and Sichuan, but not for the diversification of the lineages leading to the extant pop- ulations. Third, in spite of the large body size and rela- tive ease of sampling, Trichalophus is among the least un- derstood genera of the Holarctic weevils. Such neglect is partly due to the abundance of ambiguous historical species names, particularly in Central Asia, which creates a nomenclatorial impediment and hinders further research. Fourth, relationships of Trichalophus in Alophini, and the overall phylogenetic validity of this tribe and most of its genera (particularly Plinthodes, Pseudobarynotus mdXer- alophus) remain untested. When adequately studied, how- ever, the genus Trichalophus is expected to be of signif- icant biogeographic potential, similarly to other clades of low-dispersing organisms most suitable to reveal the ge- ographical component on their evolutionary past (Muri- enne et al. 2014; Tanzler et al. 2014; Toussaint et al. 2015). Acknowledgements. Curators of the collections mentioned above variously helped in accessing specimens under their care. Antoine Mantilleri (Paris, France) (1.) took syntype habitus and label images of both Fairmaire’s names (Fig. 5); (2.) critically verified the match between both original descriptions and the im- aged syntypes (Fig. 5) to help establish their name-bearing sta- tus and (3.) took images of Barynotus laticeps syntype (present- ly Pseudobarynotus laticeps) and of Seidlitzia marrocana syn- types (Figs 8 and 7A, B, respectively). Genrikh Ed. Davidian and Boris A. Korotyaev searched on my request for the ZIN spec- imen ofXeralophus cretaceus, which was imaged (Figs 7G, H.) by Andrey V. Frolov (all St. Petersburg, Russia). Thierry Deuve Bonn zoological Bulletin 64 (2): 59-76 (Paris, France) advised on collecting sites of Father Delavay in Yunnan. David J. Clarke (Chicago, USA) collected and made available the sequenced specimen of Trichalophus alternatus (#2968). Ignacio Ribera (Barcelona, Spain) advised on imple- mentation of DNA analytical techniques. 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Russ- ian Entomological Journal 21: 57-72 Bonn zoological Bulletin 64 (2): 59-76 ©ZFMK Bonn zoological Bulletin 64 (2): 77-106 March 2016 An annotated checklist of the inland fishes of Sulawesi Friedrich Wilhelm Miesen**, Fabian Droppelmann’, Sebastian Hiillen*, Renny Knrnia Hadiaty^ & Fabian Herder' 'Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Uchthyology Laboratory, Division of Zoology, Research Center for Biology, Indonesian Institute of Science (LIPI), Cibinong, Indonesia; E-mail: fw.miesen@googlemail.com; +49 (0)228 9122 431 Abstract. Sulawesi is the largest island of the Wallacea. Here, we present an annotated checklist of fish species record- ed in Sulawesi’s inland waters. We recognize a total of 226 species from 112 genera and 56 families. Gobiidae (41 species), Adrianichthyidae (20 species) and Telmatherinidae (19 species) are most species-rich, making up a total of 43% of the total species diversity. 65 species are endemic to Sulawesi’s freshwaters, including 1 9 Tematherinidae, 1 7 Adrianichthyi- dae, and 17 Zenarchopteridae. 44% of the inland fish fauna are obligate freshwater fishes, followed by euryhaline (38%) and amphi-, ana- or diadromous (29%) taxa. 65 species have been recorded from lacustrine environments. However, we stress that the data available are not representative for the island’s freshwater habitats. The fish species diversity of the spectacular lakes is largely explored, but the riverine ichthyofaunas are in clear need of further systematic exploration. Keywords. Sulawesi, freshwater, fishes, endemism, Wallacea, SE-Asia INTRODUCTION Sulawesi is the largest island of the Wallacea, a biodiver- sity hotspot located between the Sunda and Sahul shelves (Mokodongan & Yamahira 2015, Myers et al. 2000, Whit- ten et al. 2002). The onset of the scientific investigation of Sulawesi's inland waters and its fishes dates back to the late 19^^ century (Abendanon 1915a, b, Bleeker 1855a, 1858a, b, Boulenger 1897). Exploration of species diver- sity was, and still is, in the focus of ichthyological research on the island (e.g. Hadiaty 2007, Hadiaty & Wirjoatmod- jo 2003, Hadiaty et al. 2004, Kottelat 1989a, b, c, 1990a, b, c, d, 1991, Larson 2001, Parent! 2008, 2011, Weber 1909, 1913), complemented more recently by studies on evolutionary biology (e.g. Herder & Schliewen 2010). Sulawesi’s freshwater environments are home to sev- eral endemic animal radiations that include gastropods, crustaceans, and fishes (e.g. de Bruyn et al. 2013, Herder et al. 2006a, Mokodongan & Yamahira 2015, Parent! 2011, Parent! et al. 2013, Rintelen et al. 2007a, b, Rintelen et al. 2012, Tweedley et al. 2013). Much of this diversity is restricted to species flocks confined to a few ancient lakes, systems that serve as models for the study of speciation processes (Herder & Schliewen 2010, Rintelen et al. 2010, 2012, Vaillant et al. 2011). Phylogeographic studies of the island’s freshwater animals accordingly focused mostly on lineages of molluscs, shrimps, crabs, and fishes, with fo- cus on the lake radiations (e.g. Mokodongan & Yamahi- ra 2015, Rintelen et al. 2007b, 2014, but see also de Bruyn et al. 2012, 2013; reviewed by Rintelen et al. 2012). Received: 30.10.2015 Accepted: 09.12.2015 Parent! recognized 57 species of freshwater fishes as en- demic to Sulawesi, with the majority being restricted to the ancient lakes (Parent! 2011). Most of the non-endem- ic fish species are classified as secondary or peripheral freshwater fishes (Berra 2001), with occasional records of marine species (Kottelat 1990a, Tweedley et al. 2013). Sulawesi's lakes include some of the oldest lakes on earth. Lake Poso and the Malili Lakes in the highlands of Central Sulawesi are so-called ancient lakes, exception- ally long-lived lakes that have existed for more than 100,000 years (Brooks 1950, Rintelen et al. 2012). The Malili Lakes system is known for its endemic species flock of sailfin silversides (Telmatherinidae) (Herder et al. 2006a), but also includes small radiations of ricefishes (Oryzias) and gobies {Mugilogobius, Glossogobius; Hoese et al. 2015, Kottelat 1990d, Larson et al. 2014). Moreover, the lakes and their surroundings harbour en- demic species of half beaks (Dermogetys,Nomorhamphus; Huylebrouck et al. 2012, Meisner 2001). Ricefishes {Oryzias, Adrianichthys) are the dominant fish radiation of Lake Poso (Kottelat 1990b). Remote lakes Lindu and Tiu are substantially smaller than the ancient lakes, and harbour two (Lindu) or one (Tiu) endemic (rice-) fish species (Mokodongan et al. 2014, Parent! 2008). Lake Tondano on the northern tip of North Sulawesi is the on- ly known habitat of Tondanichthys kottelati (Collette 1995), an endemic genus and species of viviparous half- beak (Collette 1995). Lake Tempe and Lake Sidereng are Corresponding editor: P Wagner 78 Friedrich Wilhelm Miesen et al. LakeTondano Lake Limboto Lake Lindu Lake Matano - Lake Mahalona Lake Towuti .-Tenggara Lake Sidenreng Lake Tern pe 150 km Fig. 1. Map of Sulawesi showing the island's administrative partition as referred to in this study, major lakes are highlighted. Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK An annotated checklist of the inland fishes of Sulawesi 79 shallow lakes in southern Sulawesi, but little remains known about their fish fauna, and its present status apart from its use in aquaculture (Hadijah et al. 2014, Tamsil 2000) (for the location of Sulawesi's lakes see Fig. 1). In contrast to the ancient lakes, Sulawesi's riverine fish fauna has attracted far less scientific interest, and the num- ber of studies is limited (de Bruyn et al. 2013, Mokodon- gan & Yamahira 2015, Schwarzer et al. 2008, Tweedley et al. 2013). A typical feature of Sulawesi's riverine en- vironments is the absence of large, slow rivers and drainages (Kottelat 1990a), in contrast to the hydrology of most other larger Indonesian islands (Stelbrink et al. 2012). The majority of drainages are rather small and short, typically with medium to high stream velocity (Kot- telat 1990a). Sulawesi's freshwaters have been subject to massive fish species introduction and alien species invasion (Herder et al. 2012a, Kottelat & Whitten 1996, Parenti 2011, Whit- ten et al. 1987). Alien fishes were introduced for food pro- duction (e.g. Oreochromis spp., Channa spp.) (Whitten et al. 1987), pest control (Poecilia reticulata), or aquarium trade (e.g. flowerhorn cichlids) (Herder et al. 2012a). The spread of populations of alien fish species in freshwater systems of the island is apparently rapid, and may pose as a threat to the native communities (e.g. Herder et al. 2012a, Tweedley et al. 2013). Purpose of this paper The literature on freshwater fish species of Sulawesi is complex and dispersed. The most recent comprehensive source covering the island’s ichthyofauna dates back to 1993 (Kottelat et al. 1993), but is not focussed on the is- land’s fishes, and meanwhile partially out-dated. Kotte- lat’s recent catalogue on “The Fishes of the Inland Wa- ters of Southeast Asia” (Kottelat 2013) includes Sulawe- si, but focuses on the nomenclature of the whole South- east Asian ichthyofauna, and the related bibliography. The present paper aims at summarizing ichthyological records from Sulawesi’s inland waters. It provides an account of actual species records, and species that have not actually been recorded, but are likely to be present according to their known distribution. It is understood that the authors do not claim that this list is complete with respect to all records ever made, but aim at providing a baseline for analysing species records, required for upcoming inves- tigations of the island’s fauna. MATERIAL AND METHODS Literature records are compiled from the scientific liter- ature; sources or records considered questionable were not included. Material examined was mostly collected during various field campaigns of the senior author's group in Su- lawesi, since 2002. Field methods applied during field- work include beach seining, dipnetting, gillnetting, scubadiving and electrofishing. Samples were either fixed in formalin (4%) prior to storage in ethanol (80%), or fixed and stored directly in pure ethanol (-98%). Specimens were determined to the lowest feasible taxonomical lev- el using the most recent literature available. The system- atic division largely follows Kottelat (2013). ZFMK: Fish collection of Zoologisches Forschungsmuseum Alexander Koenig Bonn, Germany. Coordinates with reference to ZFMK specimens are own species records, linked to the respective voucher. Records that are not linked to vouch- ers, represent visual records (F.H.); ZMH: Zoologisches Museum Hamburg in Hamburg, Germany; MZB: Muse- um Zoologicum Bogoriense in Cibinong, Indonesia. Species occurrence is classified to: Euryhaline: species with a broad tolerance towards salinity and thus can be found in marine, brackish and freshwater environments (Hiroi & McCormick 2012); anadromous: species with adults entering marine environments and reproduction in freshwaters (Daverat et al. 2012); catadromous: species that migrate into marine environments for reproduction (Daverat et al. 2012); amphidromous: species that migrate between marine and freshwater environments for purpos- es other than reproduction (Daverat et al. 2012); freshwa- ter: primary or secondary/obligate freshwater species with no marine stadium or life history phase (Berra 2001); in- troduced: non-native, introduced species. Species expected to occur in Sulawesi, but lacking ac- tual records from the island, are included as “potential”, and justified. Clearly non-native species are classified as “introduced”. Synonyms listed are restricted to the most common ones, with emphasis on publications dealing with Sulawesi’s ichthyofauna (see Kottelat 2013 for nomencla- ture). RESULTS CHONDRICHTHYES CARCHARHINIFORMES Carcharhinidae Requiem sharks: Marine; all oceans (Nelson 2006); enter brackish and freshwaters, including lakes (Grace 2001, Nelson 2006). Carcharhinus leucas (Muller & Henle, 1839) Carcharias leucas Muller & Henle, 1839 Potential: Euryhaline; worldwide in subtropical and trop- ical coastal waters; enter brackish and freshwaters (Com- pagno 1984, Heupel & Simpfendorfer 2008); no actual records for Sulawesi. Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK 80 Friedrich Wilhelm Miesen et al. PRISTIFORMES Pristidae Sawfishes: Euryhaline; enter brackish and freshwaters; distributed in all tropical and subtropical oceans (Nelson 2006, Wueringer et al. 2009). Pristis pristis (Linnaeus, 1758) Squalus pristis hmmQm, 1758 Potential: Euryhaline; worldwide in subtropical and trop- ical coastal waters; enter brackish and freshwaters; no ac- tual records for Sulawesi (Einnaeus 1758, McEachran & Carvalho 2002). MYLIOBATIFORMES Dasyatidae Stingrays: Marine; distributed throughout the Atlantic and Indo-Pacific; some species enter brackish and freshwaters (Compagno & Roberts 1982, Nelson 2006). Himantura leoparda Manjaji-Matsumoto & Last, 2008 Potential: Euryhaline; enter brackish and freshwaters; no actual records for Sulawesi; distributed throughout the In- do-West-Pacific (Manjaji-Matsumoto & East 2008). Himantura uarnak (Gmelin, 1789) Raja uarnak GmQlm, 1789 Potential: Euryhaline; enter brackish and freshwaters; no actual records for Sulawesi; distributed throughout the In- do-Pacific (Compagno et al. 1989, Gmelin 1789). Himantura undulata (Bleeker, 1852) Trygon undulata Bleeker, 1 852d Potential: Euryhaline; enter brackish and freshwaters; no actual records for Sulawesi; distributed throughout the In- do-West Pacific (East & Stevens 1994). Himantura tutul Borsa, Durand, Shen, Alyza, Solihin & Berrebi, 2013 Potential: Euryhaline; enter brackish and freshwaters; dis- tributed throughout the Indo Pacific; no actual records from Sulawesi (Borsa et al. 2013). ACTINOPTERYGII ELOPIFORMES Megalopidae Tarpons: Euryhaline; enter brackish and freshwaters; dis- tributed in tropical and subtropical regions (Adams et al. 2013, Nelson 2006). Megalops cyprinoides (Broussonet, 1782) Clupea cyprinoides Broussonet, 1782 Euryhaline; enter brackish and freshwaters; record from Badjoa, Sulawesi Selatan (Adams et al. 2013, Bleeker 1865a, Coates 1987). ALBULIFORMES Albulidae Bonefishes: Euryhaline; enter brackish and freshwaters; distributed throughout tropical regions (Adams et al. 2013, Nelson 2006). Albula glossodonta (Forskal, 1775) Argentina glossodonta Forskal, 1775 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Myers 1991, Randall & Bauchot 1999). ANGUILLIFORMES Anguillidae Freshwater eels: Catadromous; adults inhabit freshwaters or estuaries; marine reproduction; juveniles enter fresh- waters after metamorphosis; distributed throughout trop- ical and subtropical regions except the South Atlantic and Eastern Pacific (Arai et al. 1999, Nelson 2006). Anguilla bicolor McClelland, 1844 Catadromous; distributed throughout the Indo-Pacific (Arai et al. 1999, Kottelat 2013); record from Buton (Tweedley et al. 2013). Anguilla celebesensis Kaup, 1857 Anguilla ancestralis Ege, 1939 Catadromous; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Arai et al. 1999); recorded from Eake Tondano, Manado, Sulawesi Utara (Ege 1939), Buton and Kabaena (Tweedley et al. 2013), Takes of Gorontalo (Haryono & Tjakrawidjaja 2004), Sulawe- si Utara (Arai et al. 2003), Sulawesi Tengah (Arai et al. 2003). Anguilla interioris Whitley, 1938 Potential: Catadromous; enter brackish and freshwaters; distributed throughout the Indo-Pacific; no actual records for Sulawesi (Arai et al. 1999, Kottelat 2013). Anguilla marmorata Quoy & Gaimard, 1824 Catadromous; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Arai et al. 1999); records from Sulawesi Utara (Haryono et al. 2002), Sulawesi Tenggara (02°56.035’S 121°06.855’E, ZFMK 066057), Sulawesi Selatan (3°41.589’S 119°38.629’E, ZFMK 69560), Eake Poso, Sulawesi Tengah (visual record F.H.). Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK An annotated checklist of the inland fishes of Sulawesi 81 Anguilla nebulosa McClelland, 1844 Catadromous; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Arai et al. 1999); record from Sulawesi Barat (2°37.368S 1 19°08.784’E, ZFMK 69701). Anguilla obscura Gunther, 1872 Potential: Catadromus; distributed throughout the Pacif- ic; no actual records for Sulawesi (Arai et al. 1999, Giin- ther 1872a). Moringuidae Worm, Spaghetti eels: Euryhaline; enter brackish and freshwaters; fossorial lifestyle; distributed throughout the tropical Western Atlantic and the Indo-Pacific (Nelson 2006, Tsukamoto et al. 2014). Moringua guthriana (McClelland, 1844) Ptyobranchus arundinaceus McClelland, 1 844 Potential: Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Kottelat 2013); no actual records from Sulawesi. Moringua javanica (Kaup, 1856) Aphthalmichthys javanicus Kaup, 1856 Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Allen & Steene 1988); record from Ba- ton (Tweedley et al. 2013). Moringua microchir Bleeker, 1853 Potential: Euryhaline; enter brackish waters; distributed throughout the Indo-West Pacific (Keith et al. 2006); no actual records for Sulawesi. Moringua raitaborua (Hamilton, 1822) Moringua latebrosa Schultz, 1953 Euryhaline; enter brackish waters (Kottelat 2013); record from: Kwandang, Gorontalo (Castle 1968, Kottelat 2013, Smith 1994). Muraenidae Moray eels: Euryhaline; enter brackish and freshwaters; worldwide distributed in all tropical regions (Nelson 2006, Tsukamoto et al. 2014). Gymnothorax polyuranodon (Bleeker, 1853) Muraena polyuranodon Bleeker, 1853f Potential: Euryhaline; enter brackish and freshwaters; dis- tributed throughout the Indo-Pacific; no actual records from Sulawesi (Ebner et al. 2011, Tsukamoto et al. 2014). Gymnothorax tile (Hamilton, 1822) Muraenophis tile Hamilton, 1 822 Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Tsukamoto et al. 2014); record from Ba- ton (Tweedley et al. 2013). Strophidon sathete (Hamilton, 1822) Muraenophis sathete Hamilton, 1 822 Muraena macrurus Bleeker, 1 854b Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific; record from Buton (Tweedley et al. 2013). Ophichthidae Snake, Worm eels: Marine; some species enter freshwa- ters; cosmopolitan, distributed throughout tropical regions (Cosker et al. 2012, Eschmeyer 2015, Froese & Pauly 2014, Nelson 2006). Cirrhimuraena chinensis Kaup, 1856 Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific; record from Makassar, Sulawesi Selatan (Kaup 1857). Lamnostoma mindora (Jordan & Richardson, 1908) Coecula mindora Jordan & Richardson, 1908 Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific; record from Buton (Tweedley et al. 2013). Muraenichthys gymnopterus Bleeker, 1852 Muraena gymnopterus Bleeker, 1 852b Muraenichthys microstomus Bleeker, 1 864 Euryhaline; enter brackish waters; distributed throughout the West-Pacific; record from Makassar, Sulawesi Sela- tan (Bleeker 1864). Ophichthus polyophthalmus Bleeker, 1864 Potential: Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Kottelat 2013); no actual records from Sulawesi. Pisodonophis cancrivorus (Richardson, 1848) Ophisurus cancrivorus Richardson, 1848 Ophisurus brachyosoma Bleeker, 1852b Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific; record from Makassar, Sulawesi Selatan (Bleeker 1852b). Yirrkala kaupii (Bleeker, 1858) Sphagebranchus kaupii Bleeker, 1858b Catadromous; enter brackish and freshwaters; distribute throughout Asia; record from Klabat Diatas, Sulawesi Utara (Bleeker 1858b). Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK 82 Friedrich Wilhelm Miesen et al. GONORHYNCHIFORMES Chanidae Milkfishes: Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Berra 2001; Nelson 2006). Chanos chanos (Forsskal, 1775) Mugil chanos 111 5 Euryhaline; enter brackish and freshwaters (Allen et al. 2002); record from Sulawesi Selatan (4°07.456’S 119’37.196’E, ZFMK 69759). CYPRINIFORMES Cyprinidae Minnows, Carps: Freshwater; native throughout Africa, Eurasia and North America, introduced worldwide; cyprinids are naturally absent from Sulawesi (Kottelat 1990a, Nelson 2006). Barbonymus gonionotus (Bleeker, 1849) Barbus gonionotus Bleeker, 1 849a Introduced: Freshwater; record from Sulawesi Selatan (3°41.589S 119°38.629’E, ZFMK 69514-69516, 69534, 69552-69555), Eake Poso, Sulawesi Tengah (Kottelat 1990b). Carassius auratus (Linnaeus, 1758) Introduced: Freshwater; common ornamental fish; breed- ing form from East Asia; distributed almost worldwide; record from the Malili Eakes system, Sulawesi Selatan (Nasution & Aisyah 2013), Eake Poso, Sulawesi Tengah (Kottelat 1990b). Cyprinus carpio (Linnaeus, 1758) Introduced: Freshwater; native to Central Asia; record from Malili Lakes system, Sulawesi Selatan; Lake Poso, Sulawesi Tengah (Kottelat 1990b); aquaculture escapees or stocked specimens (Herder et al. 2012a). Cyprinus cf. rubrofuscus (Lacepede, 1803) Cyprinus rubro-fuscus Lacepede, 1 803 Introduced: Freshwater; ornamental carp varieties; possi- bly derived from C. fuscus or hybrids (See Kottelat & Freyhof 2007, and references therein); record from Poso River, Sulawesi Tengah (visual record F.H.). Osteochilus vittatus (Valenciennes, in Cuvier und Va- lenciennes, 1842) Rohita vittata Valenciennes, in Cuvier und Valenciennes, 1842 Osteochilus hasselti (Valenciennes, in Cuvier und Valen- ciennes, 1842) Introduced: Freshwater; native to mainland Southeast Asia; records from Malili Lakes system and Lake Siden- reng, Sulawesi Selatan (Omar 2010); Lake Poso, Sulawe- si Tengah (visual record F.H.). CHARACIFORMES Characidae Characins: Freshwater; native to Central America, South America, and Africa; without native members in Asia (Es- chmeyer 2015, Froese & Pauly 2014, Nelson 2006). Colossoma macropomum (Cuvier, 1816) Myletes macropomus Cuvier, 1816 Introduced: Freshwater; native South America (Santos et al. 2007); record from Lake Matano, Sulawesi Selatan (Herder et al. 2012a) and Lake Poso, Sulawesi Tengah (vi- sual record F.H.). SILURIFORMES Plotosidae Eeltail catfishes: Euryhaline; enter brackish and freshwa- ters (Nelson 2006, Usman et al. 2013); distributed throughout the Indo-West Pacific (Eschmeyer 2015, Froese & Pauly 2014, Usman et al. 2013). Plotosus canius Hamilton, 1822 Introduced: Amphidromous; enter brackish and freshwa- ters (Usman et al. 2013); record from Sulawesi Selatan (04°07.540S 119°37.295’E, ZFMK 066013). Clariidae Airbreathing catfishes: Freshwater; native throughout Africa, Syria, South and West Asia (Nelson 2006, Ng & Kottelat 2007); 16 species ocurring in Asia (Ng & Kotte- lat 2007). Clarias batrachus (Linnaeus, 1758) Introduced: Freshwater; native to India, Indochina, Sun- daland and the Philippines; neotype from Java, Indonesia (Nelson 2006, Ng & Kottelat 2007); record from the Malili Lake system, Sulawesi Selatan (Herder et al. 2012a); all non-Java specimens are considered as related species- complex (Ng & Kottelat 2007). CLUPEIFORMES Engraulididae Anchovies: Euryhaline; enter brackish and freshwaters; distributed throughout the Atlantic and Indo-Pacific (Nelson 2006, Whitehead et al. 1988). Bonn zoological Bulletin 64 (2); 77-106 ©ZFMK An annotated checklist of the inland fishes of Sulawesi 83 Thrissina baelama (Forskal, 1775) Clupea baelama Forskal, 1775 Thryssa baelama (Forskal, 1775) Potential: Euryhaline; enter brackish waters (Whitehead et al. 1988); no actual records for Sulawesi; distributed throughout the Indo-Pacific (Kottelat 2013). Thrissina encrasicholoides (Bleeker, 1852) EngrauUs encrasicholoides Bleeker, 1851a Thriyssa encrasicholoides Bleeker, 1852a Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Whitehead et al. 1988, Wongratana et al. 1999). Thrissina mystax (Bloch, in Schneider, 1801) Stolephorus hamiltoni Bleeker, 1 872 Thryssa mystax (Bloch in Schneider, 1801) Euryhaline; enter brackish waters; distributed throughout the Indo-West Pacific (Wongratana et al. 1999); record from Badjoa, Sulawesi Selatan (Bleeker, 1872). Clupeidae Herrings, Shads, Sardines: Euryhaline; distributed world- wide (Nelson 2006, Wongratana et al. 1999). Herklotsichthys quadrimaculatus (Riippell, 1837) Clupea quadrimaculata Riippell, 1837 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Wongratana et al 1999). MUGILIFORMES Mugilidae Mullets: Euryhaline; enter brackish and freshwaters; dis- tributed throughout all tropical and temperate regions (Cu- vier & Valenciennes 1836, Durand 2012, Nelson 2006). Cestraeus plicatilis Valenciennes, in Cnvier & Valenci- ennes, 1836 Catadromous; enter brackish and freshwaters; distributed throughout the Indo-West Pacific (Harrison & Senou 1999); record from Sulawesi freshwaters (Valenciennes, in Cuvier and Valenciennes 1836). Mugil cephalus Linnaens, 1758 Catadromous; marine, enter brackish and freshwaters (Harrison & Senou 1997); record from Sulawesi Selatan (4°07.456’S 119’37.196’E, ZFMK 69741, 69760-69763). ATHERINIFORMES Telmatherinidae Sailfin silvers ides: With exception of Kalyptatherina helodes, on islands off the Vogelkop peninsula, restricted to Sulawesi; adaptive radiations in the Malili Eakes: Eakes Matano, Mahalona, Towuti, Eontoa, and connecting / sur- rounding rivers and streams (Hadiaty & Wirjoatmodjo 2002, Hadiaty et al. 2004, Herder et al. 2006a, b, Pfaen- der et al. 2011; reviewed by Herder & Schliewen (2010)). Telmatherina abendanoni Weber, 1913 Freshwater; endemic to Eake Matano, Sulawesi Selatan; predatory, benthic “sharpfin” species of Telmatherina (Ha- diaty & Wirjoatmodjo 2002, Herder et al. 2006a). Telmatherina antoniae Kottelat, 1991 Freshwater; endemic to Eake Matano, Sulawesi Selatan; a “roundfin” Telmatherina., the formal name Telmatheri- na antoniae is currently applied to two distinct popula- tions, T antoniae “small” and “large”; males in distinct colour morphs, females with cryptic polymorphism (Ha- diaty & Wirjoatmodjo 2002, Herder et al. 2006a, Herder et al. 2008, Pfaender et al. 2014). Telmatherina bond Weber & de Beaufort, 1922 Freshwater; endemic to rivers and streams of the Malili Eakes system, and adjacent systems; enter Eakes Matano, Mahalona and Towuti, Sulawesi Selatan; the only formal name currently available for stream-dwelling Telmathe- rina (Hadiaty & Wirjoatmodjo 2002, Herder et al. 2006a). Telmatherina celebensis Boulenger, 1897 Freshwater; endemic to Eakes Mahalona and Towuti, Su- lawesi Selatan; additional record from River Tominanga; common in both lakes; males in distinct colour morphs (Hadiaty et al. 2004, Herder et al. 2006a). Telmatherina cf. celebensis “Lontoa” Freshwater; endemic to Eake Eontoa [also Eantoa or Wa- wontoa], and surrounding swamps, Sulawesi Selatan; smaller and deeper bodied than T celebensis from Eakes Mahalona and Towuti, Sulawesi Selatan; males in distinct colour morphs (Herder et al. 2006a). Telmatherina “elongated” Freshwater; endemic to Eake Matano, Sulawesi Selatan; predatory, benthic “sharpfin” Telmatherina with short fins and slender body (Herder et al. 2006a). Telmatherina obscura Kottelat, 1991 Freshwater; endemic to Eake Matano, Sulawesi Selatan; small, blackish inshore “sharpfin” Telmatherina (Hadiaty & Wirjoatmodjo 2002, Herder et al. 2006a). Telmatherina opudi Kottelat, 1991 Freshwater; endemic to Eake Matano, Sulawesi Selatan; small “sharpfin” Telmatherina predominantly inhabiting well- structured shallows; males in distinct colour morphs (Hadiaty & Wirjoatmodjo 2002, Herder et al. 2006a). Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK 84 Friedrich Wilhelm Miesen et al. Telmatherina prognatha Kottelat, 1991 Freshwater; endemic to Lake Matano Sulawesi Selatan; large, slender “roundfin” Telmatherina, conspicuous “beak-like” jaws; epibenthic, predatory ecology; males in distinct colour morphs (Hadiaty & Wirjoatmodjo 2002, Herder et al. 2006a). Telmatherina sarasinorum Kottelat, 1991 Freshwater; endemic to Lake Matano, Sulawesi Selatan; benthic “sharpfm” Telmatherina specialized on eating eggs of sailfin silversides including conspecifics; males in dis- tinct colour morphs (Hadiaty & Wirjoatmodjo 2002, Cer- wenka et al. 2012, Herder et al. 2006a, Gray et al. 2007, Gray et al. 2008a, Gray et al. 2008b, Pfaender et al. 2010). Telmatherina “thicklip” Freshwater; endemic to Lake Matano, Sulawesi Selatan; benthic “sharpfin” Telmatherina specialized on eating shrimps; deep-bodied species, with pronounced “puffy lips” (Herder et al. 2006a, Pfaender et al. 2010). Telmatherina wahjui Kottelat, 1991 Freshwater; endemic to Lake Matano, Sulawesi Selatan; “sharpfm” Telmatherina occurring at the outlet of Lake Matano to River Petea, and the shallows of the lake (Ha- diaty & Wirjoatmodjo 2002, Herder et al. 2006a). Paratherina cyanea Aurich, 1935 Freshwater; endemic to Lake Towuti and Lake Mahalona, Sulawesi Selatan; slender, conspicuously large-eyed (Ha- diaty et al. 2004, Herder et al. 2006a). Paratherina labiosa Aurich, 1935 Freshwater; possibly endemic to Lake Lontoa, Sulawesi Selatan; holotype destroyed (Kottelat 1990c); Kottelat (1990c) tentatively assigned four juveniles obtained in 1989 to P. labiosa, further investigations lacking; no records during recent surveys by F.H. in Lake Lontoa. Paratherina striata Aurich, 1935 Freshwater; endemic to Lakes Towuti and Mahalona, Su- lawesi Selatan; largest sailfin silverside species; males in distinct colour morphs (Hadiaty et al. 2004, Herder et al. 2006a, Kottelat 1990c). Paratherina wolterecki Aurich, 1935 Freshwater; endemic to Lake Mahalona, Sulawesi Sela- tan; blackish male colouration (Hadiaty et al. 2004, Herder et al. 2006a). Tominanga aurea Kottelat, 1990 Freshwater: endemic to Lake Mahalona, Sulawesi Sela- tan; enter rivers; Kottelat (1990c) distinguished Tominan- ga aura from Tominanga sanguicauda by male colour traits, gill raker counts, and occurrence (Lake Mahalona vs. Lake Towuti); Herder et al. (2006a) reported less clear indications for species discrimination based on colour traits, and highlight the need for more detailed examina- tions (Kottelat 1990c, Hadiaty et al. 2004, Herder et al. 2006a). Tominanga sanguicauda Kottelat, 1990 Freshwater; endemic to Lake Towuti, Sulawesi Selatan; enter rivers; see Tominanga aurea for notes on species dis- crimination (Hadiaty et al. 2004, Kottelat 1990c). Marosatherina ladigesi (Ahl, 1936) Telmatherina ladigesi Ahl, 1936 Freshwater; endemic to the Bantimurung area, Maros karst, Sulawesi Selatan; dwelling in cool karst streams; males with conspicuously elongated blackish rays in sec- ond dorsal and anal fins; popular aquarium species (Ha- diaty 2007); locally transferred for breeding purposes (F.H., pers. obs.). Phallostethidae Priapium fishes: Euryhaline; enter brackish and freshwa- ters; distributed in Southeast Asia; distinct reproductive morphology: males transfer sperm with a conspicuous pri- aprium, on the underside of the head (Parenti 1996). Neostethus djajaorum Parenti & Louie, 1998 Euryhaline; endemic to Sulawesi Selatan; brackish waters of coastal plains (Parenti & Eouie 1998). BELONIFORMES Adrianichthyidae Ricefishes: Brackish and freshwaters; distributed through- out the West-Pacific; 17 of the 35 species recognized are endemic to Sulawesi, including species flocks in Eake Poso and the Malili Eakes system, endemics in small, re- mote lakes {Oryzias hadiatyae, O. soerotoi), one riverine pelvic brooder O. eversi, and a riverine lineage species from Southeast Sulawesi (Herder et al. 2012b, Kottelat 1990d, Mokodongan & Yamahira 2015, Parenti 2008, Par- enti & Hadiaty 2010, Parenti et al. 2013). Adrianichthys kruyti Weber, 1913 Adrianichthys kruytii (Soeroto & Tungka, 1991) Freshwater; endemic to Eake Poso, Sulawesi Tengah; pelagic; only a few specimens known (Kottelat 1990b, Parenti 2008). Adrianichthys oophorus (Kottelat, 1990) Xenopoecilus oophorus Kottelat, 1990a Freshwater; endemic to Eake Poso, Sulawesi Tengah; pelagic pelvic brooder; abundant in open waters of the lake Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK An annotated checklist of the inland fishes of Sulawesi 85 in 1995 (Parent! 2008); confirmed by own fieldwork in 2013; caught in the night at the surface; exploited by sub- sistence fisheries (F.H., pers. obs.). Adrianichthys poptae (Weber & de Beaufort, 1922) Xenopoecilus poptae Weber & de Beaufort, 1 922 Freshwater; endemic to Lake Poso, Sulawesi Tengah; pelagic, rare; not recorded by the authors (Kottelat 1990a, Mokodongan & Yamahira 2015, Parent! 2008, Parent! & Soeroto 2004, Soeroto & Tungka 1991, 1996). Adrianichthys roseni Parenti & Soeroto, 2004 Adrianichthys kruyti (Weber, 1913) Freshwater; endemic to Lake Poso, Sulawesi Tengah; known from a single collection; likely pelagic; abdomi- nal concavity points towards pelvic brooding (Parenti & Soeroto 2004). Oryzias asinua Parenti, Hadiaty, Lumbantobing & Herder, 2013 Freshwater; endemic to Sulawesi Tenggara; known only from the type locality: Asinua River, regency of Kendari (Parenti et al. 2013). Oryzias bonneorum Parenti, 2008 Xenopoecilus sarasinorum (Rosen, 1964) Freshwater; endemic to Lake Lindu, Sulawesi Tengah; probably pelagic (Parenti 2008). Oryzias celeb ensis (Weber, 1894) Haplochilus celebensis Weber, 1 894b Aplocheilus celebensis Weber & de Beaufort, 1912 Freshwater; records from rivers, streams and Lake Tempe in Sulawesi Selatan (Herder & Chapuis 2010, Parenti 2008), and East Timor (see Parenti 2008). Oryzias eversi Herder, Hadiaty & Nolte, 2012 Freshwater; endemic; reported only from the type local- ity in Sulawei Selatan, Tana Toraja; Salo Sadang drainage, close to village Tilanga, about 8 km south of Rantepao; the only known riverine Adrianichthyid with pelvic-brood- ing reproduction (Herder et al. 2012b). Oryzias hadiatyae Herder & Chapuis, 2010 Freshwater; endemic to Lake Masapi, Malili Lakes (Larona) system, Sulawesi Selatan; Lake Masapi is a small and shallow blackwater lake in the hills west of Lake Towuti (Herder & Chapuis, 2010). Oryzias javanicus (Bleeker, 1854) Aplocheilus javanicus Bleeker, 1 854b Euryhyaline; enter brackish waters; distributed from Thai- land to Eombok, Borneo and Sulawesi (Parenti 2008); records from Sulawesi Barat (3°20.143S 119°10.179E, ZFMK 69890-69947). Oryzias marmoratus (Aurich, 1935) Aplocheilus marmoratus Amieb, 1935 Freshwater; endemic to Takes Towuti, Mahalona, Eontoa [also Eantoa or Wawontoa], and adjacent streams, Sulawe- si Selatan; possible hybridization among O. marmoratus and O. profundicola in Take Towuti (Herder & Chapuis 2010, Kottelat 1990d, Mokodongan & Yamahira 2015). Oryzias matan ensis (Aurich, 1935) Freshwater; endemic to Take Matano, Malili Takes sys- tem, Sulawesi Selatan; abundant around the lake; shallows to deeper habitats along the coast (Kottelat 1990d, F.H., pers. obs.). Oryzias nebulosus Parenti & Soeroto, 2004 Freshwater; endemic to Take Poso, Sulawesi Tengah (Par- enti & Soeroto, 2004); small Oryzias (up to 33 mm SE); benthopelagic; male courtship at rocky, open deeper habi- tats (F.H., pers. obs.). Oryzias nigrimas Kottelat, 1990 Freshwater; endemic to Take Poso, Sulawesi Tengah; ben- thopelagic, at open habitats in the shallows (Kottelat 1990d, Parenti & Soeroto, 2004); comparatively abundant (F.H., pers. obs.). Oryzias orthognathus Kottelat, 1990 Freshwater; endemic to Take Poso, Sulawesi Tengah (Par- enti 2008); characterized by conspicuously upwards di- rected mouth, and rounded body (Kottelat 1990d, F.H., pers. obs.); tentatively pelagic, possibly rather ben- thopelagic (Parenti 2008). Oryzias profundicula Kottelat, 1990 Freshwater; endemic to Take Towuti, Sulawesi Selatan; lacustrine, deep-bodied Oryzias with filamentous fm rays; tends to inhabit deeper inshore habitats (Kottelat 1990d, F.H., pers. obs.). Oryzias sarasinorum (Popta, 1905) Haplochilus sarasinorum Popta, 1905 Xenopoecilus sarasinorum Regan, 1911 Freshwater; endemic to Take Eindu, Sulawesi Tengah; slender, pelagic pelvic brooder (Parenti 2008); Juveniles recorded in the shallows of the lake in 2013. Oryzias soerotoi Mokodongan, Tanaka & Yamahira, 2014 Freshwater; endemic to Take Tiu in Sulawesi Tengah, a small (approx. 2 km long) blackwater lake draining to the Eaa River; subadults in structured shallows; habitat of adults unknown (Mokodongan et al. 2014). Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK Friedrich Wilhelm Miesen et al. Oryzias wolasi Parenti, Hadiaty, Lumbantobing & Herder, 2013 Freshwater; endemic; small, comparatively deep-bodied Oryzias from streams in Sulawesi Tenggara, south of Kendari (Parenti et al. 2013). Oryzias woworae Parenti & Hadiaty, 2010 Freshwater; endemic to Muna island, off Sulawesi Teng- gara (Parenti & Hadiaty 2010); small comparatively deep- bodied, with conspicuous, colourful male ornamentation (Parenti & Hadiaty 2010). Zenarchopteridae Halfbeaks: Freshwater; distributed in inland and coastal habitats of the Indo-West Pacific; four genera in Sulawe- si; Nomorhamphus and Dermogenys are viviparous, less is known about the reproductive biology of Tondanichthys and Zenarchopterus; Nomorhamphus inhabit hillstreams; 12 species endemic to Sulawesi, especially species-rich (Huylebrouck et al. 2014). Halfbeak taxonomy largely fol- lows the checklist of Collette 2004, supplemented by re- cent descriptions (Anderson & Collette 1991, Collette 1995, Grier & Collette 1987; Lovejoy et al. 2004, Meis- ner 2001) Dermogenys orientalis (Weber, 1894) Hemiramphus orientalis Weber, 1 894b Dermogenys montanus Brembach, 1982 Freshwater: endemic to a hillstream of Maros Karst, Ban- timurung, Sulawesi Selatan (Collette 2004, Parenti 2011). Dermogenys vogti Brembach, 1982 Freshwater; most likely endemic to a hillstream on Lime- stone Mountain at “Topobulu”, Sulawesi Selatan [locali- ty not confirmed] (Brembach 1982, Collette 2004, Paren- ti 2011). Nomorhamphus brembachi Vogt, 1978 Nomorhamphus ravnaki Brembach, 1991 Nomorhamphus ravnaki australe Brembach, 1991 Nomorhamphus sanussii Brembach, 1991 Freshwater; endemic to Maros highland, Sulawesi Sela- tan (Collette 2004, Parenti 2011). Nomorhamphus celebensis Weber & de Beaufort, 1922 Freshwater; endemic to Lake Poso, Sulawesi Tengah (Col- lette 2004, Parenti 2011). Nomorhamphus ebrardtii (Popta, 1912) Hemiramphus (Dermatogenus) ebrardtii Popta, 1912 Freshwater; endemic to Sulawesi Tenggara; records from Wowoni Island (ZMH 7150); Muna Island, off Sulawesi Tenggara; stream leading to Lake Towuti, Sulawesi Sela- tan; stream crossing the road Soroako to Malili, 02°38.16LS, 121°12.920’E, ZFMK 49156-49176; Maros Regency, Sungai Abbalu, Village Camba, Camba District, MZB 21295 (Huylebrouck et al. 2014). Nomorhamphus hageni (Popta, 1912) Hemiramphus hageni Popta, 1912 Freshwater; endemic to Sulawesi Tenggara, Penango and Rumbia valley (Collette 2004, Parenti 2011). Nomorhamphus kolonodalensis Meisner & Louie, 2000 Freshwater; Sulawesi Tengah, Nuha drainage, north of Lake Matano, Sulawesi Selatan, and city of Poso, district of Kolonodale, Sulawesi Tengah (Collette 2004, Meisner & Louie 2000, Parenti 2011). Nomorhamphus lanceolatus Huylebrouck, Hadiaty & Herder, 2014 Freshwater: endemic to Sungai Wawolambo, Sulawesi Tenggara (Huylebrouck et al. 2014). Nomorhamphus liemi Vogt, 1978 Nomorhamphus liemi snijdersi Vogt, 1978 Freshwater; endemic to Maros highland, Sulawesi Sela- tan (Collette 2004, Parenti 2011, Vogt 1978). Nomorhamphus megarrhamphus (Brembach, 1982) Dermogenys megarrhamphus Brembach, 1982 Freshwater; endemic to Lakes Towuti and Mahalona, Su- lawesi Selatan (Collette 2004; Parenti 2011). Nomorhamphus rex Huylebrouck, Hadiaty & Herder, 2012 Freshwater; disjunct distribution in Sulawesi; records from drainage adjacent to Malili Lakes: Stream Wewu, Cerekang drainage, west of Lake Matano; stream within Malili Lakes drainage: Toletole River at village Toletole; and Sulawesi Selatan, Tana Toraja, about 8 km south of Rantepao (Huylebrouck et al. 2012). Nomorhamphus Sagittarius Huylebrouck, Hadiaty & Herder, 2014 Freshwater: Endemic to three streams in Sulawesi Teng- gara (Huylebrouck et al. 2014). Nomorhamphus towoetii Ladiges, 1972 Normorhamphus towoetii Eadiges, 1972 Freshwater; records from Eake Towuti, Sulawesi Selatan and Eake Poso, Sulawesi Tengah (Collette 2004, Eadiges 1972, Parenti 2011). Nomorhamphus weberi (Boulenger, 1897) Hemirhamphus weberi Boulenger, 1 897 Freshwater; endemic to Eakes Matano and Mahalona, Su- lawesi Selatan (Boulenger 1897, Collette 2004, Parenti 2011 ). Bonn zoological Bulletin 64 (2); 77-106 ©ZFMK An annotated checklist of the inland fishes of Sulawesi 87 Tondanichthys kottelati Collette, 1995 Freshwater; endemic to Lake Tondano, Sulawesi Utara; monotypic genus (Collette 1995, 2004, Parent! 2011). Zenarchopterus gilli Smith, 1945 Freshwater; enter brackish and coastal waters (Donaldson & Myers 2002); record from Sulawesi Selatan (4°07.456’S 119’37.196’E, ZFMK 69726-69740, 69838). Zenarchopterus dispar (Valenciennes, in Cnvier & Va- lenciennes, 1847) Hemiramphus dispar Valenciennes, in Cuvier & Valenci- ennes, 1847 Zenarchopterus maculosus Garman, 1903 Zenarchopterus vaisiganus Jordan & Seale, 1906 Potential: Freshwater; enter brackish and coastal waters; distributed throughout the Indo-Pacific; no actual records from Sulawesi (Donaldson & Myers 2002, Garman 1903). CYPRINODONTIFORMES Aplocheilidae Rivulines: Freshwater; enter brackish waters; native in the Neotropics, Africa and Southern Asia (Eschmeyer 2015, Froese & Pauly 2014, Nelson 2006). Aplocheilus panchax (Hamilton, 1822) Esox panchax Hamilton, 1 822 Possibly introduced: Freshwater; native to India and Southeast Asia; records from Buton (Tweedley et al. 2013), Sulawesi Utara (Haryono et al. 2002), Eakes of Gorontalo (Haryono & Tjala-awidjaja2004), Sulawesi Se- latan (3°4 1.589’ S 119°38.629’E, ZFMK 69557), Sulawe- si Barat (2°39.081’S 119°12.436’E, ZFMK 69651), Eake Poso, Sulawesi Tengah (visual record F.H.) and the Malili Eakes system, Sulawesi Selatan (Herder et al. 2012a). Poeciliidae Eivebearers: Freshwater; enter brackish waters; native to North, Central and South America; several worldwide in- troduced species (Nelson 2006). Gambusia affinis (Baird & Girard, 1853) Heterandria affinis Baird & Girard, 1853 Introduced: Freshwater; native to North and Central Amer- ica; introduced into warm waters almost worldwide (Pyke 2006); record from Eake Poso, Sulawesi Tengah (visual record F.H.). Gambusia holbrooki Girard, 1859 Introduced: Freshwater; native to North and Central Amer- ica; introduced to warm waters almost worldwide; likely confused with G. affinis (Girard 1859, Pyke 2006). Poecilia reticulata Peters, 1859 Introduced: Freshwater; native to northern South Ameri- ca; introduced almost worldwide; record from Malili Eakes system, Sulawesi Selatan (Herder et al. 2012a), Eake Poso, Sulawesi Tengah (1°46.29’S 120°42.98’E, ZFMK 69801-69803). GASTEROSTEIFORMES Syngnathidae Pipefishes, Seahorses: Catadromous; enter marine, brack- ish and freshwaters; distributed throughout the Atlantic, Indo-Pacific (Nelson 2006, Wilson & Orr 2011). Belonichthys mento (Bleeker, 1856) Syngnathus mento BlQokQY, 1856a Catadromous; record from Manado, Sulawesi Utara (Bleeker 1856a), Buton (Tweedley et al. 2013). Coelonotus biocellatus Gunther, 1870 Potential: Catadromous; no actual records for Sulawesi; distributed throughout the East Indian Archipelago (Giin- ter 1870, cited in Kottelat 2013). Coelonotus leiaspis (Bleeker, 1854) Syngnathus leiaspis Bleeker, 1854c Microphis leiaspis Bleeker, 1 854c Syngnathus budi Bleeker, 1856a Catadromous; distributed throughout the Indo-Pacific (Dawson 1985); records from Manado, Sulawesi Utara (Bleeker 1856a), Buton (Tweedley et al. 2013). Doryichthys boaja (Bleeker, 1850) Syngnathus boaja Bleeker, 1 850 Doryichtys spinosus Kaup, 1856 Catadromous; distributed throughout Asia (Dawson 1985); record from Makassar, Sulawesi Selatan (Kaup 1856). Hippichthys cyanospilos (Bleeker, 1854) Potential: Catadromous; no actual records for Sulawesi; distributed throughout the Indo-Pacific (Bleeker 1 854c). Hippichthys heptagonus Bleeker, 1849 Potential: Catadromous; no actual records for Sulawesi; distributed throughout the Indo-Pacific (Bleeker 1849b). Hippichthys spicifer (Riippell, 1838) Syngnathus spicifer Kuyi^QW, 1838 Catadromous; distributed throughout the Indo-Pacific (Dawson 1985); record from Sulawesi Selatan (04°14.475’S 119°36.826’E, ZFMK 066326). Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK Friedrich Wilhelm Miesen et al. Hippocampus waleananus Gomon & Kuiter, 2009 Catadromous; endemic to the Togian islands, off Sulawe- si Tengah (Gomon & Kuiter 2009). Lophocampus retzii (Bleeker, 1856) Syngnathus retzii Bleeker, 1856a Catadromous; distributed throughout the Indo-Pacific; record from Manado, Sulawesi Utara (Bleeker 1856a). Microphis ocellatus (Duncker, 1910) Doryichthys ocellatus Duncker, 1910 Catadromus; distributed throughout the Indo-Pacific (Dawson 1984); record from Sulawesi Tengah (00°55.395’S 122°52.962’E, ZFMK 066065). Oostethus brachyurus (Bleeker, 1854) Syngnathus brachyurus Bleeker, 1 854c Syngnathus polyacanthus Bleeker, 1856a Doryichthys auronitens Kaup, 1 856 Catadromous; distributed throughout the Indo-Pacific; record from Manado, Sulawesi Utara (Bleeker 1856a), Makassar, Sulawesi Selatan (Dawson 1985, Kaup 1856). Oostethus manadensis (Bleeker, 1856) Syngnathus manadensis Bleeker, 1856a Catadromous; distributed throughout the Indo-Pacific (Dawson 1985); record from Manado, Sulawesi Utara (Bleeker 1856a). SYNBRANCHIFORMES Synbranchidae Swamp eels: Freshwater; entering brackish waters; distrib- uted throughout Central and South America, Mexico, the Indo Australian Archipelago, Asia and West Africa (Nel- son 2006, Rosen & Greenwood 1976). Monopterus albus (Zuiew, 1793) Muraena alba Zuiew, 1793 Introduced: Freshwater; enter brackish waters; records from Sulawesi Utara (Haryono et al. 2002), Malili Lake drainage, Sulawesi Selatan (Herder et al. 2012a). SCORPAENIFORMES Tetrarogidae Wasp fishes: Euryhaline; enter brackish and freshwaters; distributed throughout the Indo-West Pacific (Eschmey- er 2015, Froese & Pauly 2014 Nelson 2006). Neovespicula depressifrons (Richardson, 1848) Apistus plagiometopon Bleeker, 1853a Euryhaline; entering brackish waters; record from Bu- lukumba, Sulawesi Selatan (Bleeker 1853a). Tetraroge barbata (Cuvier, in Cuvier & Valenciennes, 1829) Apistus barbatus Cuvier, in Cuvier & Valenciennes, 1 829 Euryhaline; enter brackish and freshwaters (Fricke et al. 2011); record from Sulawesi Tengah (00°55.395’S 122°52.962’E, ZFMK 066003). Tetraroge nigra (Cuvier, in Cuvier & Valenciennes, 1829) Apistus nigra Cuvier, in Cuvier & Valenciennes, 1 829 Euryhaline; enter brackish waters; record from Buton (Tweedley et al. 2013). Platycephalidae Flatheads: Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Nelson 2006). Grammoplites scaber (Linnaeus, 1758) Cottus scaber Linnaeus, 1758 Potential: Euryhaline; no actual records for Sulawesi; dis- tributed throughout the Indo-Pacific (Knapp 1999). PERCIFORMES PERCOIDEI Ambassidae Asiatic glassfishes: Euryhaline; enter brackish and fresh- waters; distributed throughout the Indo-West Pacific (An- derson & Heemstra 2003, Nelson 2006). Ambassis gymnocephala (La Cepede, 1802) Lutjan gymnocephale La Cepede, 1 802 Ambassis dussumieri Cuvier, in Cuvier & Valenciennes, 1828 Euryhaline; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Anderson & Heemstra 2003); record from Sulawesi Tengah (00°55.395’S 122°52.962’E, ZFMK 066031-066039). Ambassis interrupta Bleeker, 1853 Euryhaline; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Anderson & Heemstra 2003); type locality in Sulawesi (Bleeker 1853c). Ambassis miops Gunther, 1872 Euryhaline; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Kottelat 2013); record from Buton (Tweedley et al. 2013). Ambassis urotaenia Bleeker, 1852 Potential: Euryhaline; enter brackish and freshwaters (An- derson & Heemstra 2003); no actual records for Sulawe- si; distributed throughout the Indo-West Pacific (Bleeker 1852c). Bonn zoological Bulletin 64 (2); 77-106 ©ZFMK An annotated checklist of the inland fishes of Sulawesi 89 Ambassis vachellii Richardson, 1846 Ambassis telkaraWhitlQy, 1935a Potential: Euryhaline; enter braekish and freshwaters; no actual records for Sulawesi; distributed throughout the In- do-Pacific (Kottelat 2013). Opistognathidae Jawfishes: Euryhaline; enter brackish waters; distributed throughout the Western and Central Atlantic and Indo-Pa- cific (Kottelat 2013; Nelson 2006, Smith- Vaniz 1999). Stalix moenensis (Popta, 1922) Gnathypops moenensisVo^tdL, 1922 Euryhaline; enter brackish waters (Smith- Vaniz 1999); record from Muna Island, off Sulawesi Tenggara (Pop- tal922). Family Carangidae Jacks, Pompanos: Marine; juveniles enter brackish waters; distributed throughout the Indo-Pacific and Atlantic (Holland et al. 1996, Nelson 2006). Caranx melampygus Cuvier, in Cuvier & Valenciennes, 1833 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Holland et al. 1996). Caranx papuensis Alleyne & Macleay, 1877 Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Holland et al. 1996); record from Ba- ton (Tweedley et al. 2013). Caranx sexfasciatus Quoy & Gaimard, 1825 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Holland et al. 1996). Scontberoides lysan (Forskal, 1775) Scomber lysan Forskal, 1775 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Smith- Vaniz 1999). Selaroides leptolepis (Cuvier, in Cuvier & Valenciennes, 1833) Caranx leptolepis Cuvier, in Cuvier & Valenciennes, 1 833 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Smith- Vaniz 1999). Leiognathidae Slimys, Slipmouths, Ponyfishes: Euryhaline; enter brack- ish and freshwaters; distributed throughout the Indo-West Pacific (Eschmeyer 2015, Froese & Pauly 2014; Nelson 2006). Eubleekeria splendens (Cuvier, 1829) Equula splendens Cuvier, 1 829 Potential: Euryhaline; no actual records for Sulawesi; dis- tributed throughout the Indo-Pacific; enter brackish wa- ters (Kottelat 2013). Lutjanidae Snappers: Euryhaline; enter brackish waters; distributed throughout the Atlantic and Indo-Pacific (Eschmeyer 2015, Froese & Pauly 2014, Nelson 2006). Lutjanus argentimaculatus (Forskal, 1775) Sciaena argentimaculata Forskal, 1775 Mesoprion taeniops Valenciennes, in Cuvier & Valenci- ennes, 1830 Euryhaline; enter brackish waters; distributed throughout the Indo-West Pacific (Anderson & Allen 1999); record from Sulawesi freshwaters (Valenciennes, in Cuvier & Va- lenciennes 1830). Lutjanus bohar (Forskal, 1775) Sciaena bohar ForskM, 1775 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-West Pacific (Anderson & Allen 1999). Lutjanus ehrenbergii (Peters, 1869) Mesoprion ehrenbergii Peters, 1 869 Lutjanus oligolepis Bleeker, 1873a Euryhaline; enter brackish waters; distributed throughout the Indo-West Pacific (Anderson & Allen 1999); record from Makassar, Sulawesi Selatan (Bleeker 1873a). Lutjanus fulviflamma (Forskal, 1775) Sciaena fulviflamma Forskal, 1775 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Anderson & Allen 1999). Lutjanus fulvus (Forster, in Schneider, 1801) Holocentrus fulvus Forster, in Schneider, 1801 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Anderson & Allen 1999). Lutjanus fuscescens (Valenciennes, in Cnvier & Valen- ciennes, 1830) Mesoprion fuscescens Valenciennes, in Cuvier & Valen- ciennes, 1830 Euryhaline; enter brackish waters; distributed throughout the Indo-West Pacific (Anderson & Allen 1999); record Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK 90 Friedrich Wilhelm Miesen et al. from Sulawesi freshwaters (Valenciennes, in Cuvier & Va- lenciennes 1830). Lutjanus maxweberi Popta, 1921 Lutianus max weberi Popta, 1921 Euryhaline; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Anderson & Allen 1999); record from Kali La River, Muna Island, off Sulawesi Tenggara (Popta 1921). Haemulidae Grunts: Euryhaline; enter brackish and freshwaters distrib- uted throughout the Atlantic and Indo-Pacific (Eschmey- er 2015, Froese & Pauly 2014). Pomadasys argenteus (Forskal, 1775) Sciaena argentea Forskal, 1775 Pristipoma manadense Gunther, 1 872b Euryhaline; enter brackish waters; distributed throughout the Indo-West Pacific (McKay 2001); record from Man- ado, Sulawesi Utara (Gunther 1872b). Nemipteridae Threadfin breams: Euryhaline; enter brackish waters; dis- tributed throughout the Indo-West-Pacific (Eschmeyer 2015, Froese & Pauly 2014, Nelson 2006). Nemipterus peronii (Valenciennes, in Cnvier & Valen- ciennes, 1830) Dentex peronii Valenciennes, in Cuvier & Valenciennes, 1830 Dentex Smithii Steindachner, 1 868 Euryhaline; enter brackish waters; distributed throughout the Indo-West Pacific (Russell 2001); record from Tiworo, Muna Island, off Sulawesi Tenggara (Steindachner 1 868). Lethrinidae Emperors, Scavengers: Euryhaline; enter brackish waters; distributed from West Africa to the Indo-West Pacific (Es- chmeyer 2015, Froese & Pauly 2014, Nelson 2006). Lethrinus nebulosus (Forskal, 1775) Sciaena nebulosaVoxs\M, Ml 5 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Carpenter 2001a). Polynemidae Threadfins: Euryhaline; enter brackish and freshwaters; distributed in all tropical and subtropical regions (Moto- mura 2004, Nelson 2006). Polydactylus microstoma (Bleeker, 1851) Polynemus microstoma Bleeker, 1851a Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Feltes 2001); record from Bulukumba, Sulawesi Selatan (Bleeker 1851a). Sciaenidae Drums: Euryhaline; enter brackish and freshwaters; Dis- tributed throughout the Atlantic and Indo-Pacific (Es- chmeyer 2015, Froese & Pauly 2014, Nelson 2006). Nibea soldado (La Cepede, 1802) Holocentrus soldado La Cepede, 1 802 Corvina celebica Bleeker, 1 854d Euryhaline; enter brackish waters; distributed throughout the Indo-West Pacific (Sasaki 2001); record from Makas- sar, Sulawesi Selatan (Bleeker 1854d). Mullidae Goatfishes: Euryhaline; enter brackish waters; distributed throughout the Atlantic and Indo-Pacific (Eschmeyer 2015, Froese & Pauly 2014; Nelson 2006). Upeneus tragula Richardson, 1846a Upeneus sundaicus yar. caudalis Popta, 1921 Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Randall 2001); record from Tiworo, Mu- na Island, off Sulawesi Tenggara (Popta 1921). Toxotidae Archerfishes: Euryhaline; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Berra 2001 , Nel- son 2006). Toxotes chatareus (Hamilton, 1822) Coins chatareus Hamilton, 1 822 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Allen 1991, 2001). Toxotes jaculatrix (Pallas, in Schlosser, 1767) Sciaena jaculatrix Pallas, in Schlosser, 1767 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Allen 1991, 2001). Terapontidae Grunters, Tigerperches: Euryhaline; enter brackish and freshwaters; distributed throughout the Indo-West Pacif- ic (Berra 2001, Nelson 2006, Vari 2001). Lagusia micracanthus (Bleeker, 1860) Datnia micracanthus Bleeker, 1860 Therapon {Datnia) micracanthus Bleeker, 1873b Terapon micracanthus Fowler, 1931 Papuservus micracanthus Munro, 1958 Euryhaline; endemic to Sulawesi; enter brackish and fresh- waters; records from Lagusi, Amparang, Bantimurung, Bonn zoological Bulletin 64 (2); 77-106 ©ZFMK An annotated checklist of the inland fishes of Sulawesi 91 Cendrana, Leang-leang, Maros, Menralang, Samanggi and Saripa rivers, Manjali Spring, Sulawesi Selatan (Bleeker 1860, Fowler 1931, Vari & Hadiaty 2012). Ter upon jarbua (Forskal, 1775) SciaenajarbuaYoxskkX, 111 5 Euryhaline; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Vari 2001); record from Su- lawesi Selatan (04°14.475’S 119°36.826’E, ZFMK 066043). Kuhliidae Flagtails: Euryhaline; enter brackish and freshwaters; dis- tributed throughout the Indo-Pacific (Berra 2001, Nelson 2006). Kuhlia marginata (Cuvier, in Cuvier and Valenciennes, 1829) Dules marginatus Cuvier, in Cuvier & Valenciennes, 1 829 Euryhaline; enter brackish and freshwaters; distributed throughout the Indo-West Pacific (Carpenter 2001b); records from Buton (Tweedley et al. 2013), Sulawesi Utara (Haryono et al. 2002), Sulawesi Selatan (3°30.822’S 119°32.267’E, ZFMK 69614-69615). Kulia rupestris (La Cepede, 1802) Centropomus rupestris Ea Cepede, 1 802 Euryhaline; enter brackish and freshwaters; distributed throughout the Indo-West Pacific (Carpenter 2001b); record from Buton (Tweedley et al. 2013). LABROIDEI Cichlidae Cichlids: Diverse group in marine, brackish and freshwa- ter environments; distributed throughout the Neo- and Palaeotropics; not native to Sulawesi; several species in- troduced worldwide (Berra 2001, Eschmeyer 2015, Froese & Pauly 2014; Nelson 2006). “Flowerhorn” cichlid Introduced: Freshwater; hybrid of neotropical species; records from the Malili Eakes system, Sulawesi Selatan (Herder et al. 2012a), Poso River, Sulawesi Tengah (vi- sual record F.H.). Melanochromis cyaneorhabdos (Bowers & Stauffer, 1997) Introduced: Freshwater; native to Eake Malawi, East Africa; record from the Malili Eakes system, Sulawesi Se- latan (Herder et al. 2012) and Eake Poso, Sulawesi Ten- gah (visual record F.H.). Oreochromis mossambicus (Peters, 1852) Chromis mossambicus Peters, 1 852 Introduced: Freshwater; native to Africa; record from the Malili Eakes system, Sulawesi Selatan (Herder et al. 2012a, Nasution & Aisyah 2013), Eake Poso, Sulawesi Tengah, and various streams (visual record F.H.). Oreochromis niloticus (Linnaeus, 1758) Introduced: Freshwater; native to Africa; record from the Malili Lakes system (Nasution & Aisyah 2013), Sulawe- si Selatan and Sulawesi Barat (2°39.08r S 1 19°12.436’E, ZFMK 69650). Scaridae Parrotfishes: Euryhaline; enter brackish waters; distributed throughout the Atlantic and Indo-Pacific (Eschmeyer 2015, Froese & Pauly 2014, Nelson 2006). Chlorurus sordidus (Forskal, 1775) Scarus sordidusVoxskkX, 1775 Scarus celebicus Bleeker, 1 854d Euryhaline; enter brackish waters (Bellwood 2001), record from Makassar, Sulawesi Selatan (Bleeker 1854d). BLENNIOIDEI Blenniidae Combooth blennies: Euryhaline; enter brackish and freshwaters; distributed throughout the Atlantic and Indo- Pacific (Eschmeyer 2015, Froese & Pauly 2014, Nelson 2006). Meiacanthus anema (Bleeker, 1852) Petroskirtes anema Bleeker, 1 852c Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Allen 1991); record from Kabaena (Tweedley et al. 2013). CALLIONYMOIDEI Callionymidae Dragonets: Euryhaline; two species in freshwaters; distrib- uted throughout the Indo-West Pacific (Eschmeyer 2015, Froese & Pauly 2014, Nelson 2006). Eleutherochir opercularis (Valenciennes, in Cuvier & Valenciennes, 1837) Callionymus opercularis Valenciennes, in Cuvier & Va- lenciennes, 1837 Brachycallionymus mirus Herre, 1936 Euryhaline; enter brackish waters; distributed throughout the Indo-West Pacific (Talwar & Jhingran 1991); record from Lembeh Strait, north coast of Sulawesi (Herre, 1936). Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK 92 Friedrich Wilhelm Miesen et al. GOBIOIDEI Rhyacichthyidae Loach gobies: Freshwater; distributed throughout the In- do-West Pacific (Berra 2001, Nelson 2006). Rhyacichthys aspro (Valenciennes, in Cnvier and Va- lenciennes, 1837) Platyptera aspro Valenciennes, in Cuvier & Valenciennes, 1837 Anadromous; enter brackish and freshwaters; distributed throughout Indo-Pacific (Allen 1991); records from Ba- ton (Tweedley et al. 2013), Sulawesi Utara (Haryono et al. 2002), Sulawesi Barat (3°16.65rS 118°51.929’E, ZFMK 6848-6850). Eleotrididae Sleepers: Euryhaline; enter brackish and freshwaters; dis- tributed worldwide in tropical and subtropical regions (Berra 2001, Eschmeyer 2015, Froese & Pauly 2014, Nel- son 2006). Belobranchus belobranchus (Valenciennes, in Cnvier & Valenciennes, 1837) Eleotris belobrancha Valenciennes, in Cuvier & Valenci- ennes, 1837 Anadromous; distributed throughout the Indo-Pacific (Allen 1991); records from Manado, Sulawesi Utara (Va- lenciennes, in Cuvier & Valenciennes, 1 837), Buton and Kabaena (Tweedley et al. 2013), Sulawesi Utara (Hary- ono et al. 2002) and Sulawesi Barat (2°39.308’S 119°12.095’E, ZFMK 69631; 2°39.08rS 119 12.436’E, ZFMK 69642-69647; 2°38.428’S 119°09.294’E, ZFMK 69670; 2°37.368’S 119°08.784’E, ZFMK 69699). Belobranchus segura Keith, Hadiaty & Lord, 2012 Freshwater; enter brackish and freshwaters; described from Halmahera and Irian Jaya (Keith et al. 2012); record from Sulawesi Barat (3°16.65US 118°51.929’E, ZFMK 69814-69815). Bostrychus microphthalmus Hoese & Kottelat, 2005 Freshwater; endemic to Gua Tanette cave, Sulawesi Se- latan; genus poorly defined and likewise poorly known (Hoese & Kottelat 2005). Bunaka gyrinoides (Bleeker, 1853) Eleotris gyrinoides Bleeker, 1 853c Anadromous; distributed throughout the Indo-West Pacif- ic (Allen 1991); records from Buton (Tweedley et al. 2013), Sulawesi Selatan (3°41.589’S 119°38.629’E, ZFMK 69556, 69558-69559), Sulawesi Barat (3° 16. 65 US 118°51.929’E, ZFMK 69823). Butis amboinensis (Bleeker, 1854) Eleotris amboinensis Bleeker, 1 854a Anadromous; enter brackish and freshwaters; distributed throughout the Indo-West Pacific (Yokoo et al. 2006); record from Buton (Tweedley et al. 2013). Butis butis (Hamilton, 1822) Cheilodipterus butis Hamilton, 1 822 Potential: Anadromous; enter braekish and freshwaters; no actual records for Sulawesi; distributed throughout the In- do-Pacific (Allen et al. 2002). Eleotris fusca (Schneider, 1801) Poecilia fusca Schneider, 1801 Potential: Anadromous; enter brackish and freshwaters as adults; no actual records for Sulawesi; distributed through- out the Indo-Pacific (Maeda et al. 2007). Eleotris melanosoma Bleeker, 1853 Anadromous; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Bleeker 1853c, Maeda et al. 2007); records from Sulawesi Selatan (3°30.822'S 119°32.267’E, ZFMK 69595, 69616-69617), Sulawesi Barat (3°16.65rS 118°51.929’E, ZFMK 69806). Giuris margaritaceus (Valenciennes, in Cuvier & Va- lenciennes, 1837) Eleotris margaritacea Valenciennes, in Cuvier & Valen- ciennes, 1837 Anadromous; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Allen et al. 2002); record from Sulawesi Barat (3°16.65rS 118°51.929’E, ZFMK 69804-69805, 69822). Oxyleotris marmorata (Bleeker, 1852) Eleotris marmorataBlQokQr, 1852e Anadromous; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Earson & Murdy 2001); record from Sulawesi Selatan (3°41.589'S 119°38.629’E, ZFMK 69512-69513, 69550-69551). Gobiidae Gobies: Euryhaline, catadromous, anadromous and fresh- water; includes a total of 1725 species in 25 1 genera (Berra 2001, Eschmeyer 2015, Froese & Pauly 2014, Nelson 2006). Acentrogobius janthinopterus (Bleeker, 1853) Gobius janthinopterus Bleeker, 1 853b Gobius hemigymnopomus Bleeker, 1856a Amphidromous; enter brackish and freshwaters; distrib- uted throughout the Indo-Pacific (Donaldson & Myers 2002); record from Makassar, Sulawesi Selatan (Bleeker 1853b). Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK An annotated checklist of the inland fishes of Sulawesi 93 Acentrogobius moloanus (Herre, 1927) Anadromous; enter brackish and freshwaters; distributed throughout the Western Pacific (Blaber & Milton 1990); record from Sulawesi Selatan (119°36.826’E 04°14.475’S, ZFMK 066064). Amblygobius decussatus (Bleeker, 1855) Gobius decussatus Bleeker, 1855a Anadromous; enter brackish and freshwaters; distributed throughout the Western Central Pacific (Myers 1991); record from Manado, Sulawesi Utara (Bleeker 1855a). Awaous grammepomus (Bleeker, 1849) Gobius grammepomus Bleeker, 1 849c Anadromous; enter brackish and freshwaters; distributed throughout Asia (Watson 1992); records from Sulawesi Selatan (3°41.589'S 119°38.629’E, ZFMK 69502-69511, 69535-69549; 3°30.822’S 119°32.267’E, ZFMK 69602- 69603, 69628), Sulawesi Barat(2°38.428'S 119°09.294’E, ZFMK 69653-69659; 2°37.915'S 119°09.488’E, ZFMK 69673-69676). Bathy gobius petrophilus (Bleeker, 1853) Gobius petrophilus Bleeker, 1853d Gobius villosus Weber, 1909 Anadromous; entering brackish and freshwaters; distrib- uted throughout the Indo-West Pacific; record from Man- ado, Sulawesi Utara (Weber 1 909, Weber & de Beaufort 1953). Cryptocentroides insignis (Seale, 1910) Amblyogobius insignis Seale, 1910 Cryptocentroides dentatus Popta, 1922 Anadromous; entering brackish and freshwaters; distrib- uted throughout the Western Pacific; record from Raha, Muna Island, off Sulawesi Tenggara (Popta 1922). Drombus bontii (Bleeker, 1849) Gobius bontii Bleeker, 1 849c Acentrogobius elberti Popta, 1921 Anadromous; entering brackish and freshwaters; distrib- uted throughout the Indo-West Pacific (Kottelat 2013); record from Raha, Muna Island, off Sulawesi Tenggara (Popta 1921). Glossogobius celebius (Valenciennes, in Cnvier & Va- lenciennes, 1837) Freshwater; endemic to Sulawesi; records from Eake Towuti, Sulawesi Selatan (119°37.295’E 04°07.540’S, ZFMK 066014); Sulawesi Selatan (3°41.589'S 119°38.629’E, ZFMK 69517; 3°30.822’S 119°32.267’E, ZFMK 69596-69601, 69612-69613, 69626-69627), Su- lawesi Barat (2°37.368’S 119°08.784’E, ZFMK 69697- 69698). Bonn zoological Bulletin 64 (2): 77-106 Glossogobius flavipinnis (Anrich, 1938) Freshwater; endemic to Eake Towuti, Sulawesi Selatan; lacustrine dwarf species (Hoese et al. 2015). Glossogobius intermedius (Anrich, 1938) Freshwater; endemic to Eakes Mahalona and Towuti, Su- lawesi Selatan (Kottelat et al. 1993). Glossogobius matanensis (Weber, 1913) Freshwater; endemic to Eakes Matano, Mahalona, Towu- ti, and Eontoa, Sulawesi Selatan (Kottelat et al. 1993). Glossogobius mahalonensis Hoese, Hadiaty & Herder, 2015 Freshwater; endemic to Eake Mahalona, Sulawesi Sela- tan; so far known from one single site within the lake (Hoese et al. 2015). Gnatholepis anjerensis (Bleeker, 1851) Gobius anjerensis Bleeker, 1851b Anadromous; enter brackish waters; distributed through- out the Indo-Pacific; record from Bunaken Island, off Manado, Sulawesi Utara (Bleeker, 1851b). Lentipes mekonggaensis Keith, Hadiaty, Hubert, Bns- son & Lord, 2014 Presumably amphidromous; terra typica is a fast flowing stream in Sulawesi Tenggara (Keith et al. 2014). Lentipes watsoni Allen, 1997 Presumably anadromous; record from Sulawesi Tenggara (02°65.035'S 121°06.855'E, ZFMK 45041); type locali- ty in Papua New Guinea; determination based on one male specimen matching the diagnosis of the species descrip- tion (Allen 1997). Lophogobius bleekeri Popta, 1921 Anadromous; enter brackish and freshwaters; distributed throughout the Western Pacific; recorded from Raha, Mu- na Island, off Sulawesi Tenggara (Popta 1921). Mugilogobius adeia Larson & Kottelat, 1992 Freshwater; endemic to Lake Matano, Malili Lakes sys- tem, Sulawesi Selatan (Larson 2001, Larson & Kottelat 1992). Mugilogobius amadi (Weber, 1913) Gobius amadi Weber, 1913 Weberogobius amadi Koumans, 1953 Freshwater; endemic to Lake Poso, Sulawesi Tengah; orig- inally abundant, not reported since 1985 (Larson 2001). Mugilogobius chulae (Smith, 1932) Vaimosa chulae Smith, 1932 Freshwater; distributed throughout the South-East Asian ©ZFMK 94 Friedrich Wilhelm Miesen et al. archipelago, southern Japan, Sri Lanka, Thailand, Malaysia, Taiwan and Singapore (Huang et al. 2013, Tan & Lim, 2004); record from Boloang, Sulawesi Utara (Lar- son 2001, Larson et al. 2008). Mugilogobius hitam Larson, Geiger, Hadiaty & Her- der, 2014 Freshwater; most likely endemic to Lake Towuti, Sulawe- si Selatan (Larson et al. 2014). Mugilogobius latifrons (Boulenger, 1897) Gobius latifrons Boulenger, 1 897 Gobius latifrons WQhQX, 1913 Vaimosa latifrons Aurich, 1938 Freshwater; endemic to streams and lakes of the Malili Lakes system, Sulawesi Selatan (Larson et al. 2014). Mugilogobius lepidotus Larson, 2001 Freshwater; endemic to Lake Towuti, Malili Lakes sys- tem, Sulawesi Selatan; recorded by F.H. in 2002 and 2004. Mugilogobius mertoni (Weber, 1911) Gobius mertoni Wober, 1911 Tamanka mindora Herre, 1945 Vaimosa layia Herre, 1953 Tamanka mertoni Koumans, 1953 Potential: Anadromous; distributed throughout the Indo- Pacific; no actual records for Sulawesi (Heemstra et al. 2004, Huang et al. 2013, Larson 2001, Larson et al. 2013, Manilo & Bogorodsky 2003). Mugilogobius rexi Larson, 2001 Freshwater; endemic to Lake Mahalona and Lake Towu- ti, Sulawesi Selatan (Larson 2001). Mugilogobius sarasinorum (Bonlenger, 1897) Gobius sarasinorum Boulenger, 1 897 Tamanka sarasinorum Koumans, 1953 Freshwater; endemic to Lake Poso, Sulawesi Tengah; highly abundant in 2013 (F.H. pers. obs.). Oxyurichthys tentacularis (Valenciennes, in Cnvier & Valenciennes, 1837) Gobius tentacularis Valenciennes, in Cuvier & Valenci- ennes, 1837 Oxyurichthys rumbia Popta, 1922 Anadromous; enters brackish and freshwaters; distributed throughout the Indo-West Pacific (Mauge 1986); record from Rumbia, Sulawesi Tenggara (Popta 1922). Periophthalmus kalolo Lesson, 1831 Periophthalmus argentilineatus Valenciennes, in Cuvier & Valenciennes, 1 837 Anadromous; enter brackish and freshwaters; records from Buton (Tweedley et al. 2013), Sulawesi Selatan (04°14.475'S 119°36.826'E, ZFMK 066001-066002). Redigobius bikolanus (Herre, 1927) Vaimosa bikolana Herre, 1927 Anadromous; enter brackish and freshwaters; distributed throughout Borneo, Sulawesi and the Philippines (Kotte- lat et al. 1993); record from Buton (Tweedley et al. 2013). Redigobius penango (Popta, 1922) Pseudogobius penango Popta, 1922 Anadromous; enter brackish and freshwaters; distributed in Indonesia; record from Penango, Sulawesi Tenggara (Larson 2010, Popta 1922). Schismatogobius bruynisi de Beaufort, 1912 Anadromous; enter brackish and freshwaters; distributed throughout Indonesia and the Philippines (de Beaufort 1912, Keith & Lord 2011, Kottelat & Whitten 1996, Kot- telat et al. 1993); record from Sulawesi Tengah (121°06.855’E 02°56.035’S, ZFMK 45049). Schismatogobius marmoratus (Peters, 1868) Gobiosoma marmorataVQtQVS, 1868 Anadromous; enter brackish and freshwaters; distributed throughout Sulawesi, Philippines and Japan (Keith & Eord 2011, Kottelat et al. 1993); record from Sulawesi Tengah (02°56.035’S 121°06.855’E, ZFMK 066059; 00°55.395’S 122°52.962’E, ZFMK 066060). Sicyopterus cynocephalus (Valenciennes, in Cuvier & Valenciennes, 1837) Sicydium cynocephalum Valenciennes, in Cuvier & Valen- ciennes, 1837 Anadromous; enter brackish and freshwaters; distributed throughout Indonesia and the Philippines (Keith & Eord 2011, Koumans 1953); records from Manado harbour, Su- lawesi Utara (Valenciennes, in Cuvier & Valenciennes 1837), Sulawesi Utara (Haryono et al. 2002) and Buton (Tweedley et al. 2013). Sicyopterus longifilis de Beaufort, 1912 Anadromous; enter brackish and freshwaters (Allen 1991, Keith & Eord 2011); distributed throughout Seram, Sulawesi, Sumatra and the Philippines (Koumans 1953); records from Sulawesi Utara (Haryono et al. 2002), Su- lawesi Selatan (3°27.242’S 119°32.357’E, ZFMK 69563- 69573, 69575-69577; 3°30.822’S 119°32.267’E, ZFMK 69604-69609, 69618-69619), Sulawesi Barat(2°39.08 US 119°12.436’E, ZFMK 69632-69641; 2°38.428’S 119 09.294’E, ZFMK 69667-69673; 2°37.915’S 119°09.488’E, ZFMK 69677-69678; 2°37.368’S 119°08.784’E, ZFMK 69689-69696; 3°16.65LS 118°5L929’E, ZFMK 69816-69821, 69837). Bonn zoological Bulletin 64 (2); 77-106 ©ZFMK An annotated checklist of the inland fishes of Sulawesi 95 Sicyopterus macrostetholepis (Bleeker, 1853) Sicydium macrostetholepis Bleeker, 1853e Sicydium gymnauchen Bleeker, 1858a Anadromous; enter brackish and freshwaters; distributed throughout Indonesia and the Philippines ( Allen 1991, 2011, Keith & Lord 2011); record from Manado, Sulawe- si Utara (Bleeker 1858a), Buton and Kabaena (Tweedley et al. 2013), Sulawesi Utara (Haryono et al. 2002). Sicyopterus microcephalus (Bleeker, 1855) Sicydium microcephalus Bleeker, 1855b Anadromous; enter brackish and freshwaters; distributed throughout Asia (Allen 1991, Keith & Lord 2011); record from Buton (Tweedley et al. 2013). Sicyopterus micrurus (Bleeker, 1854) Sicydium micrurus Bleeker, 1854a Anadromous; enter brackish and freshwaters; distributed throughout Asia (Allen 1991, Keith & Lord 2011); records from Buton and Kabaena (Tweedley et al. 2013). Sicyopus zosterophorus (Bleeker, 1856) Sicydium zosterophorum Bleeker, 1856b Anadromous; enter brackish and freshwaters; distributed throughout Asia (Allen 1991); record from Sulawesi Barat (3°16.65LS 118°5L929’E, ZFMK 69810-69813, 69835- 69836). Stenogobius ophthalmoporus (Bleeker, 1854) Gobius ophthalmoporus Bleeker, 1 854a Chonophorus lachrymosus Weber, 1 894a Anadromous; enter brackish and freshwaters; distributed throughout Asia; records from Sulawesi Selatan (Watson 1991, Weber 1894a) and Buton (Tweedley et al. 2013). Stiphodon elegans (Steindachner, 1879) Anadromous; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Watson 1995); records from Buton and Kabaena (Tweedley et al. 2013), Sulawesi Ten- gah(02°56.035’S 121°06.855’E, ZFMK 066027-066028), Sulawesi Selatan (3°27.242’S 119°32.357’E, ZFMK 69578-69584; 3°30.822’S 119°32.267’E, ZFMK 69620- 69622), Sulawesi Barat (2°39.08LS 119°12.436’E, ZFMK 69648-69649; 2°38.428'S 119°09.294’E, ZFMK 69667-69669; 2°37.915’S 119°09.488’E, ZFMK 69702- 69704; 3° 1 6.65 1 ’ S 1 1 8°5 1 .929’E, ZFMK 69832-69834); specimens recorded outside the Society, Tubuai and Samoa Islands are considered as closely related to S. elegans (Kot- telat 2013). Stiphodon semoni Weber, 1895 Anadromous; enter brackish and freshwaters; distributed throughout the Indo-Pacific (Watson 1996); records from Buton (Tweedley et al. 2013), Sulawesi Utara; (Haryono et al. 2002). Yongeichthys nebulosus (Forskal, 1775) Gobius nebulosusYoxskkX, 111 5 Acentrogobius nebulosus (Forsskal, 1775) Potential: Anadromous; enter brackish and freshwaters; no actual records for Sulawesi; distributed throughout the In- do-Pacific (Randall et al. 1990). ACANTHUROIDEI Scatophagidae Scats: Euryhaline; enter brackish waters; distributed throughout the Indo-West Pacific (Berra 2001, Eschmey- er 2015, Froese & Pauly 2014, Nelson 2006). Scatophagus argus (Linnaeus, 1766) Chaetodon argus Einnaeus, 1766 Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Allen 1984); record from Buton (Tweed- ley et al. 2013). SCOMBROIDEI Sphyraenidae Barracudas: Marine; enter brackish waters; distributed in all tropical and subtropical parts of the Atlantic, Indian and Pacific Ocean (Eschmeyer 2015, Froese & Pauly 2014, Nelson 2006). Sphyraena barracuda (Edwards, in Catesby, 1771) Esox barracuda Edwards, in Catesby, 1771 Euryhaline; juveniles enter brackish waters (Kottelat 2013; Senou 2001); record from Sulawesi Selatan (4°07.456’S 1 19’37. 196’E, ZFMK 69758); original description is a re- jected work and not available as a source, author of the species description follows Kottelat (2013). Sphyraena obtusata Cuvier, in Cuvier & Valenciennes, 1829 Potential: Euryhaline; enter brackish waters; no actual records for Sulawesi; distributed throughout the Indo-Pa- cific (Senou 2001). ANABANTOIDEI Anabantidae Climbing gouramies: Freshwater; enter brackish waters; distributed throughout the Indo-West Pacific (Berra 2001 , Eschmeyer 2015, Froese & Pauly 2014, Nelson 2006). Anabas testudineus (Bloch, 1792) Anthias testudineus Bloch, 1792 Anabas variegatus Bleeker, 1851a Introduced: Freshwater; enter brackish waters; native to India, South and Southeast Asia; record from Kema, Su- lawesi Utara (Bleeker 1851a), Malili Eakes system (Na- Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK 96 Friedrich Wilhelm Miesen et al. Table 1. Summary of species and records according to i) region and ii) salt tolerance / ecology. Total numbers of records and species are highlighted, numbers of introduced and endemic species refer to these total numbers. Species with no actual record for Sulawesi are listed as "potential". Note that obligate and primary freshwater fishes are combined as freshwater fishes. The islands of Buton and Kabaena, off Sulawesi Tenggara, are treated as individual regions (Tweedley et al. 2013). sution & Aisyah 2013), Sulawesi Selatan, Lake Poso, Su- lawesi Tengah (Kottelat 1990b, confirmed in 2012 by F.H.). Osphronemidae Gouramies: Freshwater; distributed throughout Sumatra, Java and Borneo; able to breath atmospheric air using a specialised respiratory organ (Berra 2001, Kottelat et al. 1993, Nelson 2006). Trichopodus (Trichogaster) pectoralis Regan, 1910 Introduced: Freshwater; native to Thailand; record from the Malili Lakes system, Sulawesi Selatan (Herder et al. 2012a, Kottelat et al. 1993, ). Trichopodus (Trichogaster) trichopterus Pallas, 1770 Introduced: Freshwater; native to Sundaland and Indochi- na; record from the Malili Lakes system, Sulawesi Sela- tan (Herder et al. 2012a, Kottelat et al. 1993), Lake Poso, Sulawesi Tengah (Kottelat 1990b, visual record F.H.). Channidae Snakeheads: Freshwater; native to tropical Africa and Southern Asia (Berra 2001; Nelson 2006). Channa lucius (Cuvier, in Cuvier and Valenciennes, 1831) Ophicephalus lucius Cuvier, in Cuvier and Valenciennes, 1831 Introduced: Freshwater; native to Sundaland and Indochi- na; record from the Malili Lakes system, Sulawesi Sela- tan (Kottelat et al. 1993). Channa striata (Bloch, 1793) Ophicephalus striatus Bloch, 1793 Introduced: Freshwater; native to India, China and South- east Asia; records from the Malili Lake system, Sulawe- si Selatan (Hadiaty & Wirjoatmodjo 2002, Hadiaty et al. 2004, Herder et al. 2012a), Lake Poso, Sulawesi Tengah (3°4L589’S 119°38.629’E, ZFMK 69518; 3°30.822’S 119°32.267’E, ZFMK 69625) and Sulawesi Barat (2°38.428’S 119°09.294’E, ZFMK 69671-69672). PLEURONECTIFORMES Paralichthyidae Sand flounders: Euryhaline, enter brackish and freshwa- ters; distributed throughout the Atlantic and Indo-Pacific (Froese & Pauly 2014, Eschmeyer 2015, Nelson 2006). Pseudorhombus malayanus Bleeker, 1865 Euryhaline; enter brackish waters (Amaoka & Hensley 2001), record from Makassar, Sulawesi Selatan, (Bleek- er 1865a). Pseudorhombus neglectus Bleeker, 1865 Euryhaline; enter brackish waters (Amaoka & Hensley 2001); record from Makassar, Sulawesi Selatan (Bleek- er, 1865a). Soleidae Soles: Euryhaline; enter brackish and freshwaters; distrib- uted throughout tropical and temperate regions; usually flat, bottom dwelling fishes (Froese & Pauly 2014, Nel- son 2006). Bonn zoological Bulletin 64 (2); 77-106 ©ZFMK An annotated checklist of the inland fishes of Sulawesi 97 Achirus poropterus (Bleeker, 1851) Euryhaline; enter brackish and freshwaters (Bleeker 1851c); record from Sulawesi Selatan (4°07.456’S 119’37.196’E, ZFMK 69766-69767). TETRAODONTIFORMES Triacanthidae Triplespines: Euryhaline; enter brackish waters; distrib- uted throughout the Indo-Pacific (Nelson 2006, Santini & Tyler 2002). Triacanthus biaculeatus (Bloch, 1786) Balistes biaculeatus Bloch, 1786 Triacanthus russellii Bleeker, 185 Id Euryhaline; enter brackish waters (Matsuura 2001); record from Makassar, Sulawesi Selatan (Bleeker 1851d). Tetraodontidae Puffers: Euryhaline; enter brackish and freshwaters; dis- tributed throughout all tropical and subtropical parts of the Atlantic and Indo-Pacific (Nelson 2006, Yamanoue et al. 2011 ). Arothron manilensis (Marion de Proce, 1822) Tetrodon Manilensis Marion de Proce, 1 822 Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Randall 1985); record from Sulawesi Se- latan (04°07.540’S 119°37.295’E, ZFMK 066046- 066052). Chelonodontops patoca (Hamilton, 1822) Tetrodon patoca Hamilton, 1 822 Chelonodon patoca (Hamilton, 1 822) Euryhaline; enter brackish waters; distributed throughout the Indo-Pacific (Talwar & Jhingran 1991); record from Sulawesi Selatan (04°07.540’S 119°37.295’E, ZFMK 060060). Dichotomy ctere erythrotaenia (Bleeker, 1853) Tetraodon erythrotaenia Bleeker, 1853e Euryhaline; enter brackish and freshwaters; distributed throughout the Indo-West Pacific (Allen, 1991); record from Maros, Sulawesi Selatan (Bleeker 1853e). DISCUSSION Actual records and likely occurrences of fishes in inland waters of Sulawesi sum up to a total of 226 species (see Table 1 for details). This ichthyofauna is composed of 1 12 genera and 56 families, dominated by Gobiidae (41 species, 18%), Adrianichthyidae (20 species, 9%), Tel- matherinidae (19 species, 8%), and Zenarchopteridae (17 species, 7%). Taken together, these four families account for 43% of the island’s total inland fish species diversity. Sulawesi's native inland ichthyofauna is heterogeneous in terms of salt tolerance: Only 89 species (44% of all na- tive species) are obligate freshwater fishes, whereas 77 species (38% of all native species) are euryhaline. 60 species (29% of all native species) are amphi-, ana- or catadromous, migrating between marine and freshwater environments. 65 species (32% of all native species) of the species inventory are endemic. 46 (71% of all endem- ic species) of these endemic species are from radiations in the ancient lakes of Sulawesi while only 1 8 riverine in- cluding three euryhaline species are considered endemic. In addition, endemism also appears to be unevenly dis- tributed among the families. Telmatherinidae (19 species), Adrianichthyidae (17 species) and Zenarchopteridae (17 species) contain in sum 86% of all endemic Sulawesi fish- es. In 2011, Parenti reported a total of 76 native freshwa- ter fish species from Sulawesi, of which 56 were consid- ered endemic. This significantly higher number reported here arises to a smaller proportion from additional, recent species descriptions (e.g. Hoese et al. 2015, Huylebrouck et al. 2012, 2014, Earson et al. 2014, Mokodongan et al. 2014, Parenti et al. 2013). However, it is largely due to the wider focus of the present list, which includes all fish species recorded from Sulawesi's inland waters, also wide- spread fish species that are very likely to be expected in the island’s inland waters, but without actual records. Sulawesi’s freshwater and coastal habitats are facing substantial and manifold threats from habitat degradation (e.g. urbanization, damming, surface mining), and stock- ing with alien fish species (African cichlids, Asian carps, snakeheads, or gouramis, and others - see Kottelat et al. 1993, Herder, et al. 2012, Tweedley et al. 2013). The list presented here includes 22 introduced species, some of which have been recognized as potential threats to the in- digenous fauna of Sulawesi's ancient lakes (see Herder et al. 2012a for alien fish species recorded in the Malili Fakes area). The total number of fish species of Sulawesi’s freshwa- ter and brackish habitats is significantly lower than that of the Sundaic islands, like closeby Borneo with its at least 430 fish reported species (Kottelat et al. 1993, McGinley & Hogan 2003, Tan 2006; but note that no actual check- list of the inland fishes is available). However, the actu- al size of Sulawesi is four times smaller than that of Bor- Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK 98 Friedrich Wilhelm Miesen et al. neo (Rachman et al. 201 5). With values between 37% (160 species, McGinley & Hogan 2003) to 62% (267 species, Tan 2006) Bomeos rate of endemism is exceptionally high, however the four times smaller island of Sulawesi with its endemism rate of 32% (65 species), appears surpris- ingly close to its larger neighbour. The actual state of exploration of the inland ichthyofau- na of Sulawesi shows clear regional sampling biases. However, it appears clear that the total species account of fishes occurring in Sulawesi’s inland waters is strong- ly dominated by the lake species flocks (see also Tweed- ley et al. 2013, Parent! 2011, Herder & Schliewen 2010), but the exploration of the riverine fish species diversity, and its distribution across the island, remains in a gener- ally fragmentary stage. Acknowledgements. We thank the Indonesian Institute of Sci- ences (LIPI), and the State Ministry of Research and Technol- ogy (RISTEK) for the permit to conduct research in Indonesia. Fieldwork benefited from help by B. Stelbrink, J. Pfaender, S. Chapuis and M. Milanovic. F.H. thanks A.W. Nolte for fruitful ricefish discussions and joint fieldwork. We acknowledge T. von Rintelen for providing access to digitized maps. Fieldwork was funded by a research grant of the DFG (to F. Herder; DFG HE 5707/2-1), and by the ARCBC 2001-2003 project to S. Wirjoat- modjo. REFERENCES Abendanon EC (1915a) Midden-Celebes-Expeditie. Geologis- che en geographische Doorkruising van Midden-Celebes (1909-1910), Atlas. Brill, Eeiden Abendanon EC (1915b) Midden-Celebes-Expeditie. Geologis- che en geographische Doorkruising van Midden-Celebes (1909-1910) 2. Brill, Eeiden Adams AJ, Horodysky AZ, McBride RS, Guindon K, Shenker J, MacDonald TC, Harwell HD, Ward R, Carpenter K (2013) Global conservation status and research needs for tarpons (Megalopidae), ladyfishes (Elopidae) and bonefishes (Albu- lidae). Fish and Fisheries 15: 280-311. 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Ma- thematisch-Naturwissenschaftliche Classe 57; 965-1008 Steindachner F (1879) Ichthyologische Beitrage (VIII). Sitzungs- berichte der Kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Classe 80: 119-191 Stelbrink B, Albrecht C, Hall R, Rintelen T von (2012) The bio- geography of Sulawesi revisited: is there evidence for a vic- ariant origin of taxa on Wallace’s “anomalous island”? Evo- lution 66: 2252-2271 Talwar PK, Jhingran AG (1991) Inland fishes of India and ad- jacent countries. Volume 2. A.A. Balkema, Rotterdam Tamsil A (2000) Studi Beberapa Karakteristik Reproduksi Prapemijahan dan Kemungkinan Pemijahan Buatan Ikan Bun- go {Glossogohius cf aureus) di Danau Tempe dan Danau Sidenreng Sulawesi Selatan. Program Pascasarjana Institut Pertanian Bogor Tan HH (2006) The Borneo Suckers. Natural history Publica- tions (Borneo), Kinabalu Tan HH, Fim KKP (2004) Inland fishes from the Anambas and Natuna islands. South China Sea. With description of a new species of Betta (Teleostei: Osphronemidae). Raffles Bulletin of Zoology Supplement 11: 107-115 Tsukamoto K, Watanabe S, Kuroki M, Aoyama J, Miller MJ (2014) Freshwater habitat use by a moray eel species, Gym- ©ZFMK 106 Friedrich Wilhelm Miesen et al. nothorax polyuranodon, in Fiji shown by otolith microchem- istry. Environmental Biology of Fishes 97: 1377-1385. DOI 10. 1007/s 1064 1-0 14-0228-9 Tweedley JR, Bird DJ, Potter IC, Gill HS, Miller PJ, O’Dono- van G, TJakrawidJaJa AH (2013) Species compositions and ecology of the riverine ichthyofaunas in two Sulawesian is- lands in the biodiversity hotspot of Wallacea. Journal of fish biology 82: 1916-50. doi: 10.1111/jfb.l2121 Usman BI, Amin SMN, Arshad AA, Aminur Rahman M (2013) Review of Some Biological Aspects and Fisheries of Grey- Eel Catfish Plotosius canius (Hamilton, 1822). 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Zoologische Ergebnisse einer Reise inNiederlandisch Ost-Indien 3: 405-476 Weber M (1894b) Die Siisswasser-Fische des Indischen Archi- pels, nebst Bemerkungen iiber den Ursprung der Fauna von Celebes. Zoologische Ergebnisse einer Reise in Niederlan- disch Ost-Indien: 405-476 Weber M (1895) Fische von Ambon, Java, Thursday Island, dem Burnett-Fluss und von der Sud-Kuste von Neu-Guinea. In: Zoologische Forschungsreisen in Australien und dem ma- layischen Archipel; mit Unterstutzung des Herrn Dr. Paul von Ritter ausgeftihrt ...Jahren 1891-1893 von Dr. Richard Semon 5: 259-276 Weber M (1909) Diagnosen neuer Fische der Siboga-Expediti- on. Notes from the Eeyden Museum 31: 143-169 Weber M (1911) Die Fische der Aru- und Kei-Inseln. Ein Bei- trag zur Zoolographie dieser Inseln. Abhandlungen der Sen- ckenbergischen Naturforschenden Gesellschaft Frankfurt am Mam 34: 1-49 Weber M (1913) Die Fische der Siboga-Expedition. Siboga- Ex- peditie. Brill EJ, Eeiden Weber M, de Beaufort EF (1912) Over de zoetwatervisshen van Timor en Babber. Verslagen van de gewone vergaderingen der Wissen Natuurkundige Afdeeling der Koninklijke Akademie van Wetenschappen te Amsterdam 1912: 133-138 Weber M, de Beaufort, EF (1922) The fishes of the Indo-Aus- tralian Archipelago. IV. Heteromi, Solenichthyes, Synentog- nathi, Percesoces, Eabyrinthici, Microcyprini. Brill, Eeiden Weber M, de Beaufort EF (1953) The fishes of the Indo-Aus- tralian archipelago Vol. X Gobioidea. Brill, Eeiden Whitehead PJP, Nelson GJ, Wongratana T (1988) FAO Species Catalogue. Vol. 7. Clupeoid fishes of the world (Suborder Clu- peoidei). An annotated and illustrated catalogue of the her- rings, sardines, pilchards, sprats, shads, anchovies and wolf- herrings. FAO, Rome Whitley GP (1935a) Fishes from Princess Charlotte Bay, North Queensland. Records of the South Australian Museum 5: 345-365 Whitley GP (1938) Descriptions of some New Guinea fishes. 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Reviews in Fish Biology and Fisheries 19: 445-464 Yamanoue Y, Miya M, Doi H, Mabuchi K, Sakai H, & Nishida M (2011) Multiple invasions into freshwater by pufferfishes (Teleostei: Tetraodontidae): A mitogenomic perspective. PEoS One 6 (2): el 74 10 Yokoo T, Kanou K, Moteki M, Hiroshi K, Tongnunui P, Kuroku- ra H (2006) Juvenile morphology and occurrence patterns of three Butis species (Gobioidei: Eleotridae) in a mangrove es- tuary, southern Thailand. Ichthyological Research 53: 330-336 Zuiew B (1793) Biga Mvraenarvm, novae species descriptae. Nova Acta Academiae Scientiarum Imperialis Petropolitanae 7 (for 1789): 296-301 Bonn zoological Bulletin 64 (2): 77-106 ©ZFMK Bonn zoological Bulletin 64 (2): 107-108 March 2016 Correction & update On the Linck collection and specimens of snakes figured by Johann Jakob Scheuchzer (1735) - the oldest fluid-preserved herpetological collection in the world? Bauer, A.M. & R. Wahlgren (2013). Bonn Zoological Bulletin 62: 220-252 Aaron M. Bauer Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, U.S.A. aaron. bauer@villanova. edu; + 1-610-519-485 7 The contributions of Dr. Konstantin Wopke to the reor- ganization of the Linck collection in the Natural ienkabi- nett Waldenburg have been outlined by Bauer and Wahlgren (2013). Unfortunately, Dr. Wopke was inadver- tently misidentified in Figure 7 (p. 227) of this paper. Orig- inally identified as the figure on the left, he is, in fact, the figure on the right of the photo, wearing a white lab coat. The figure on the left of the photo is the mineralogist Al- fred Seifert. This photograph had earlier appeared, correct- ly labeled, in Budig (1999). Through the kindness of Mr. Dietrich Wopke, who has shared some biographical data with me, I am able to pro- vide some further information regarding his father. Kon- stantin Wopke was born 29 July 1905 in Crimmitschau, 17 km west of Waldenburg, as the son of Richard and He- lene (nee Brendel) Wopke. He attended school in Leipzig until the outbreak of World War I and then continued his schooling in Gotha. He studied zoology, botany, chemistry in Jena, Freiburg im Breisgau and in Leipzig. In 1930 he was promoted to the degree of Dr. phil. on the basis of his dissertation “Die Kloake und die Begattungsorgane der mannlichen Zauneidechse {Lacerta agilis L.)” complet- ed in Leipzig under the direction of Prof Dr. Johannes Meisenheimer (1873-1933), a specialist on the develop- ment of invertebrates, and published in Jena (Wopke 1930). He subsequently worked as a teaching assistant at the Zoological Institute of the University of Leipzig and as a research assistant at the Anatomical Institute in Wurzburg. From 29 May 1933 to 6 July 1935 he re-or- ganized the approximately 5000 zoological objects in the Furstlich-Schonburg-Waldenburgische Naturalienkabi- nett in Waldenburg, struggling with misidentifications, mislabeling, and a century or more of the intermixing of more recently acquired specimens with the 17* and 18* century material of the original Linck collection. Received: 23.03.2015 Accepted: 06.11.2015 In January 1934 Wopke passed the state examination for secondary school teachers in the subjects of zoology, botany and chemistry. After graduating from his intern- ship year at the Furstlich-Schonburgischen Deutschen Oberschule he was employed as a private tutor at the Kam- mergut in Hardisleben in Thuringia. He was subsequent- ly excluded from school employment by the Saxon Min- istry of Education on political grounds and instead, from 1936 to 1939 he served as a research assistant in the Im- perial Biological Institute in Naumburg/Saale, where he worked on “Phylloxera in the wine growing areas of Saale and Unstrut.” The documentation and results of this work, although apparently unpublished, are maintained in the German Federal Archives. Although he had moved on from Waldenburg, in 1937 Wopke’s opinion was solicit- ed about the renovation of the collections of the Franck- esche Stiftungen in Haale (Mojsejenko 2013) and in the same year his guide to the Naturalienkabinett in Walden- burg was published (Wopke 1937). In August 1939 he was sent to military service and in April 1944 he was killed in Russia. Although Wopke’s scientific career was cut short, his contribution to the reorganization of the Linck collection in Waldenburg is a lasting legacy. REFERENCES Budig U (1999) Ein Kabinett voher Raritaten - Die Einck Sammlung. Pp. 16-31 in: Budig U, Zenker R & Stadt- lander G. (eds) Heimatmuseum und Naturalienkabinett Waldenburg, SachsischeMuseen7. Sachsische Eandes- stehe fur Museumswesen, Chemnitz Mojsejenko, A (2013) Die Kunst- und Naturalienkammer der Franckeschen Stiftung in Halle an der Saale. Sem- inararbeit im Erganzungsmodul des Masterstudien- ganges, Universitat Duisburg-Essen, 24 pp. Corresponding editor: Ph. Wagner 108 Aaron M. Bauer Wopke K (1930) Die Kloake und die Begattungsorgane der mannlichen Zauneidechse (Lacerta agilis L.). Je- naische Zeitschrift fiir Naturwissenschaft 65 : 275-3 1 8, pis iv-vi Wopke K (1937) Fiihrer durch das Fiirstlich Schonburgi- sche Natural ienkabinett in Waldenburg/Sa. und die in ihm enthaltenen Sammlungen. Fiirstlich Schonburgis- che Naturalienkab inert in Waldenburg, Waldenburg, 38 pp. Bonn zoological Bulletin 64 (2); 107-108 ©ZFMK Bonn zoological Bulletin 64 (2): 109-116 March 2016 A new genus and new species of Neotropical Thoracophorini (Coleoptera: Staphylinidae: Osoriinae) Ulrich Irmler Institut for Ecosystem Research, Dept. Applied Ecology, University of Kiel, Olshausenstrasse 40, D-24098 Kiel, Germany; E-mail: uirmler@ecology. uni-kiel.de Abstract. Three new species of the new genus Geotrochopsis are described: G. pubescens, G. collaris, and G.flaveolus. The genus is placed in the subtribe Clavilispinina of the tribe Thoracophorini. G. pubescens seems to be distributed all over the Neotropical region. Both G. collaris and G.flaveolus are from Peru and Brazil, respectively. Additionally, one new species is described from the Central Amazon: Geomitopsis amazonensis. Furthermore, Ashnaosorius Makhan, 2008 is recognised as a new synonym of Geomitopsis. Key words. New species, Osoriinae, Thoracophorini, Neotropics INTRODUCTION During the studies on the Neotropical Osoriinae few spec- imens of blind species were found that belong to the tribe Thoracophorini. Hitherto among this subtribe, only the blind genera Geomitopsis Scheerpeltz, 1931 and Ash- naosorius Makhan, 2008 were known from the Neotrop- ical region. A more detailed study came to the result that some specimens belong to a new genus. The present pa- per describes this new genus with its new species. Fur- thermore, Ashnaosorius is a new synonym of Geomitop- sis. According to Herman (2001) the genus Geomitopsis is recorded also from the Mediterranean region from Libanon to Canary Islands with nine species and from Africa with six species. Including the new Geomitopsis species, a total of six species is also known from the Neotropics. Therefore, a key to the species of the Neotropical region is provided. MATERIAL AND METHODS The material studied in this investigation is presently de- posited in the following museums and private collections. AMNH American Museum of Natural History, New York BMNH British Museum, Natural History, London INPA Collections of Instiuto National de Pesquisas da Amazonia, Manaus, Brazil KNHM Kansas Natural History Museum, Lawrence NHMP National Museum of Natural History, Czec Republic, Prague Received: 15.05.2014 Accepted: 18.03.2016 ZFMK Museum Alexander Koenig, Bonn, Germany JJC Private collection of Jifi Janak, Prague, Czech Republic UIC Private collection of U. Irmler, Plon, Germany The photographs were taken using a Makroskop M 420 (Wild, Herbrugg) in combination with a digital camera (LeicaECS). CombineZS (Hadley 2006) was used to op- timise depth of focus. Length was measured in the mid- dle of tagmata: head from clypeus to posterior edge, pronotum from anterior to posterior edge along midline, elytra from anterior edge of shoulders to posterior edge; width at the widest part of tagmata (head width includes eyes). In the measurement of total length, the abdominal inter-segmental space is subtracted. The aedeagus was dis- sected and drawings were made using a microscope un- der 250 X magnification. DESCRIPTION OF THE NEW SPECIES Geotrochopsis n. gen. Type species. Geotrochopsis pubescens n.sp. is here des- ignated as the type species Diagnosis. Geotrochopsis is similar to the other blind genus in the tribe Thoracophorini, i.e. Geomitopsis Scheerpeltz, 193 1. In contrast to Geomitopsis Scheerpeltz, 1931, Geotrochopsis is densely pubescent on the whole body and the antennomere six is not narrower than anten- nomeres five and seven. Moreover, the tarsi of Geotro- chopsis are composed of five tarsomeres, whereas they are Corresponding editor: D. Ahrens 110 Ulrich Irmler composed of four tarsomeres in Geomitopsis . Addition- ally, the aedeagus of Geotrochopsis is symmetric and with- out a ventral prominence such as in Geomitopsis. Description. Length and habitus: elongate; blind with re- duced elytra; small species of about 1.3-1. 6 mm total length. Head approximately square; clypeus semicircular; labrum divided into two lobes separated by a deep emar- gination; eyes absent; setiferous punctation; without dis- crete neck; gular sutures combined. Antennae not geniculate; width of antennomeres in- creasing from second antennomere to apex of antenna; penultimate antennomere wider than long. Pronotum wider than long; sides smoothly rounded; lat- eral margin fine; with dense setiferous punctation; in pos- terior half with longitudinal medial impression. Elytra much wider than long; not longer than pronotum; shoulders widely rounded; divergent from shoulders to posterior angles; hind wings reduced; sutural striae weak or absent; with setiferous punctation. Abdomen elongate; conically narrowed posteriad; densely pubescent; with microsculpture. Protibia slightly wider than meso- and metatibia; pro- coxae slightly elongate; tarsi composed of five tarsomeres. Aedeagus with broad and stout central lobe; central lobe nearly straight; not or weakly curved; paramera as long as or longer than central lobe: spermatheca oval with short and straight ductus. Etymology. The specific name is a combination of the Greek words geo meaning earth or soil, trochus meaning circlet, and opsis standing for appearance. The name refers to the similarity to the gmus Allotrochus Fagel, 1955, and to the soil dwelling life. Geotrochopsis pubescens n. sp. Figs 2A, F, 5A, D Type material. Holotype, male: Peru . Huanuco, Pangua- na (74°56’W, 9°37’S), rain forest, collected by pitfall trap, April 1984, leg. M. Verhaagh (UIC). Paratypes: Mexico. 1 female, Veracruz, Cordoba, Para- je Nueve Nacimento, tropical evergreen forest, collected by Berlese method, 7.8.1969, leg. S. & J. Peck (AMNH); Costa Rica. 1 male, 2 females, Vulcan Arenal, rd. to Are- nal Observ. Fodge (84°43.58’W, 10°26.5rN), forest bor- der, litter & dead wood, sifted, 3.12.2012, leg. M. Schiilke (UIC, MSC); 3 females, Puntarenas, OS A Peninsula, 5km W Rincon de OSA (83°3rW, 8°42’N), 50 m elevation, forest floor, collected by Berlese method, 24.-30.3. 1973, leg. J. Wagner & J. Kethley (AMNH); 1 male, Osa Penn., Fundacion Neotrop., 10 km W. Rincon (83°31.30’W, 8°42.30’N), 20 m elevation, collected from forest litter by berlese, 23.6. 1997, leg. R. Anderson (KNHM); Peru. 1 fe- male, Rio Tambopata Reseve, 30 km SW Puerto Maldon- ado (69°16’W, 12°12’S), trop. Moist forest, on fungi Rigi- doporus microporus, 19.9.-10.10.1984, leg. D.A. Grimal- di (AMNH); Brazil. 1 male, Sao Paulo, leg. Mraz (NMP). Diagnosis. The species resembles Geotrochopsis flaveo- lus n.sp. (see below) in the structure of the aedeagus and the widely rounded posterior angles of the pronotum. It can be differentiated from G. flaveolus by the shorter in- ner lateral lobes and the longer paramera. The pronotum of G. flaveolus is more strongly arched than that of G pu- bescens, in particular, in the posterior half Description. Fength: 1.4 mm. Colouration: yellow, legs and antennae light yellow. Head: 0. 19 mm long, 0.29 mm wide; without eyes; tem- ples behind base of antennae approximately as long as clypeus; sides of temples parallel; clypeus semicircular; extremely finely punctate; pubescent; weak netlike mi- crosculpture; surface moderately shiny; small area at base of antennae without punctation and microsculpture; sur- face shiny. Antennae as long as head and pronotum combined; first antennomere thick and as long as second and third com- bined; second antennomere nearly as thick as first, but shorter; third antennomere narrower than preceding and following antennomeres; nearly half as long as second an- tennomere; following antennomeres increasing in width; penultimate antennomere nearly twice as wide as long. Pronotum: 0.23 mm long, 0.36 mm wide; widest in mid- dle; evenly narrowed in smooth curve to anterior and pos- terior margin; anterior angles obtuse; anterior edge not margined; posterior angles widely rounded; posterior edge widely emarginate and distinctly margined; in dorsal as- pect, lateral margin fine in anterior half, widened in pos- terior half and continuous in equal width to wide poste- rior margin; in central posterior half with finely impressed furrow; punctation deeper and coarser than on head; on average, interstices between punctures as wide as diam- eter of punctures; pubescent; netlike microsculpture deeper than on head; meshes on average wider than di- ameter of punctures; surface moderately shiny. Elytra: 0.22 mm long, 0.38 mm wide; widest at poste- rior angles; shoulders evenly rounded; with slight obtuse angle, only; sides distinctly margined; posterior edge of elytra straight; punctation slightly finer than on pronotum; irregular microsculpture deeper than on pronotum; pubes- cent; surface matt; less shiny than on pronotum. Abdomen as densely and finely punctate as elytra; punc- tures partly granulate; pubescent; deep netlike mi- crosculpture; surface as matt as surface of elytra. Aedeagus short and broad; apical lobe shortly curved; paramera distinctly projecting apical lobe; lateral inner lobes transverse and only slightly projecting apical lobe. Bonn zoological Bulletin 64 (2): 109-116 ©ZFMK A new genus and new species of Neotropical Thoracophorini 111 Fig l-4.Geotrochopsis jlaveolus (1), G. pubescens (2), G. collaris (3), and Geomitopsis amazonensis (4); Aedeagus in ventral and lateral aspect (A); protibia and tarsi (B); mesotibia and tarsi (C), metatibia and tarsi (D), spermatheca (E), antenna (F); scale bar: 0 . 1 . Etymology. The specific name pubescens derived from Geotrochopsis collaris n. sp. the same Latin name and means pubescent. It refers to the Figs 3A-F, 5C, E finely hairy punctation. Type material. Holotype, male: Peru. Madre de Dios, Cuzco Amazonica (69°02.06’W, 12°36.48’S), 300 m el- evation, secondary forest Wl, F95420, 17.5.1995, leg. D. Agosti (AMNH). Bonn zoological Bulletin 64 (2): 109-116 ©ZFMK 112 Ulrich Irmler Fig. 5. Dorsal aspect of head, pronotum, and elytra of Geotrochopsis pubescens (A), G. flaveolus (B), G collaris (C), lateral aspect of G pubescens (D), and ventral aspect of G collaris (E); dorsal aspect of head, pronotum, and elytra of Geomitopsis ama- zonensis (F), and G. campanae (G), G remilleti (H), lateral aspect of Geomitopsis amazonensis (I); scale bar: 0.2 mm. Bonn zoological Bulletin 64 (2): 109-116 ©ZFMK A new genus and new species of Neotropical Thoracophorini 113 Paratypes: 4 males, 16 females with same data as for holotype; 2 females, same location as holotype, but 17.5.1994 (AMNH,UIC). Diagnosis. The species can be easily differentiated from the other two species by the specific structure of the prono- tum. The pronotal sides are slightly emarginate in front of the posterior angles, whereas they are smoothly round- ed in the two other species. Furthermore, the posterior edge of the elytra is slightly retreated at suture as trian- gular emargination and the aedeagus has not the pair of inner lobes as found in G. pubescens and G flaveolus. Description. Length: 1.35 mm. Colouration: dark yellow; antennae and legs lighter yellow. Head: 0. 19 mm long, 0.26 mm wide; without eyes; tem- ples as long as semi-circular clypeus; punctation fine; pu- bescent; on average, interstices between punctures twice as wide as diameter of punctures; netlike microsculpture moderately deep; meshes distinctly wider than diameter of punctures; surface moderately shiny. Antennae longer than head and pronotum combined; first antennomere thick and nearly as long as second and third antennomere combined; second antennomere slight- ly narrower than first, but thicker than third antennomere; twice as long as third antennomere; following anten- nomeres increasing in width; third antennomere approx- imately square; penultimate antennomere nearly twice as wide as long. Pronotum: 0.23 mm long, 0.33 mm wide; widest in mid- dle; evenly narrowed to anterior angles in smooth curve; in front of posterior angles with slight emargination; pos- terior angles obtuse, but nearly rectangular; anterior edge not margined; lateral margin fine and continuous to pos- terior edge; margined posterior edge slightly emarginate; punctation slightly denser and deeper than on head; pu- bescent; on average, interstices between punctures slight- ly wider than diameter of punctures; netlike microsculp- ture as deep and wide as on head; surface moderately shiny. Elytra: 0.21 mm long, 0.35 mm wide; widest at poste- rior angles; narrowed to shoulders in smooth curve; in dor- sal aspect, lateral margin visible in its total length; shoul- ders obtuse without forming angles; posterior edge of ely- tra retreated to suture as wide triangular emargination; punctation as deep and dense as on pronotum; microsculp- ture slightly deeper; surface as shiny as on pronotum. Abdomen as densely and finely punctate as elytra; punc- tures partly granulate; pubescent; deep netlike mi- crosculpture; surface as matt as surface of elytra. Aedeagus slender with long central lobe; apical lobe as long as basal lobe; nearly straight; in slight obtuse angle to basal lobe; slender paramera slightly projecting central lobe; with numerous sensillae; inner lobe projecting. Etymology. The specific name derived from the Latin word collum for pronotum and refers to the specific struc- ture of the posterior angles of the pronotum. Geotrochopsis flaveolus n. sp. Figs lA, F, 5B Type material. Holotype, male: Brazil. Rio de Janeiro, Moro de Babilonia (43°10’W, 22°57’S), 100-200 m ele- vation, 25.10.2002, leg. J. Janak (INPA). Paratypes: 172 specimens with same data and from same location as holo- type (NHMP, JJC, UIC, ZFMK). Diagnosis. The species is characterised by the arched pronotum, in particular, in the posterior half In dorsal as- pect, the lateral margin is covered at posterior angles. Fur- thermore, the pair of inner lobe are longer than in G pu- bescens and the paramera are shorter. Description. Length: 1.6 mm. Colouration: Dark yellow; posterior margin of pronotum darker, light brownish; legs and antennae light yellow. Head: 0.25 mm long, 0.30 mm wide; without eyes; tem- ples as long as semi-circular clypeus; punctation moder- ately deep and dense; pubescent; on average, interstices between punctures twice as wide as diameter of punctures; netlike microsculpture moderately deep; meshes distinct- ly wider than diameter of punctures; surface moderately shiny. Antennae as long as head and pronotum combined; first antennomere long and thick; slightly shorter than second and third antennomeres combined; second antennomere slightly narrower and shorter than first antennomere; con- ical third antennomere much narrower and shorter than second antennomere; following antennomeres increasing in width; fourth antennomere approximately square; penultimate antennomere nearly twice as wide as long. Pronotum: 0.25 mm long, 0.39 mm wide; widest in mid- dle; narrowed in even and smooth curve to anterior and posterior angles; posterior angles obtuse; rounded in wide smooth curve without forming distinct angles; anterior edge without margin; lateral margin fine; continuous to posterior edge; in dorsal aspect, margin covered in ante- rior half and at posterior angles; punctation as dense and deep as on head; pubescent; netlike microsculpture mod- erately deep; meshes wider than diameter of punctures; with short impressed furrow in posterior half of midline; netlike microsculpture as deep and wide as on head; sur- face moderately shiny. Elytra: 0.24 mm long, 0.40 mm wide; widest at poste- rior angles; slightly narrowed in posterior half; more strongly narrowed in anterior half; shoulders widely rounded; posterior edge of elytra straight; in dorsal aspect, lateral margin visible throughout its total length; contin- Bonn zoological Bulletin 64 (2): 109-116 ©ZFMK 114 Ulrich Irmler ued to shoulders and ending shortly in front of scutellum; punctation deeper and denser than on pronotum, pubes- cent; netlike microsculpture slightly deeper than on prono- tum, but meshes as wide as on pronotum and wider than diameter of punctures; surface moderately shiny. Abdomen conically narrowed posteriad; densely punc- tate and pubescent; punctation still denser than on elytra; punctures partly granulate; netlike microsculpture dense and deep. Aedeagus broad; apical lobe placed in wide obtuse an- gle to basal lobe; inner lateral lobes distinctly projecting central lobe; paramera longer than central lobe, but short- er than central lobe and projecting inner lobes combined; group of three sensillae in middle of paramera. Etymology. The specific name derived from the Latin word 'Jlaveolus' meaning yellowish and refers to the over- all yellowish colouration of the species. Key to the species of Geotrochopsis n. gen. 1. Posterior angles of pronotum obtusely rounded; aedeagus with pair of inner lobes 2 - Posterior angles of pronotum nearly rectangular, pronotal sides in front of posterior angles slightly sin- uate (Fig. 5C), aedeagus with one inner lobe (Fig. 3 A) G. collaris n. sp. 2. Pronotum strongly arched; in dorsal aspect, margin at posterior angles covered (Fig. 5B), pair of inner lobes of aedeagus long, paramera not projecting in- ner lobes (Fig. lA) G. Jlaveolus n. sp - Pronotum not arched, in dorsal aspect, margin at pos- terior angles visible (Fig. 5A), pair of inner lobes of aedeagus short and transverse, paramera projecting in- ner lobes (Fig. 2 A) G. pubescens n. sp. Geomitopsis Scheerpeltz, 1931 Ashnaosorius Makhan, 2008: 1, new synonymy. Libanotyphlus Coiffait, 1954: 155. Remarks. Makhan (2008) described the new genus Ash- naosorius on the basis of two new species from Suriname that were formerly described under the genus Cubanoty- phlus Coiffait & Decou (1972). He characterised the genus by tarsi being composed of three tarsomeres and differ- ences in the aedeagal structure to Geomitopsis Scheerpeltz, 1931. However, he compared his species only with G. remilleti Orousset, 1985 that was placed in the subgenus Pseudogeomitopsis Orousset, 1983. Fortunately, he pub- lished photos of the legs. It can be derived from the pho- tos that the tarsi are not composed of three tarsomeres, as mentioned in the description, but composed of four tar- someres as in the genus Geomitopsis. The first tarsomere is very short and can be easily overlooked. Furthermore, all other characters of Geomitopsis, i.e. sixth antennomere narrower than fifth and seventh antennomeres, absence of eyes, the specific structure of the aedeagus, and the over- all habitus as derived from the photos of the original pub- lication are equal 'm Ashnaosorius and Geomitopsis. Thus, no generic difference between Ashnaosorius and Geomi- topsis is found. Therefore, Ashnaosorius is regarded as synonym to Geomitopsis. Unfortunately, the type specimens of the species of Ash- naosorius could not be studied. Requests to loan speci- mens were not answered. Geomitopsis amazonensis n. sp. Figs 4A-F, 5F Type material. Holotype, male: Brazil, Amazonas, Reserva Ducke, 26 km NE Manaus, Plot A, leaf litter, Jan. 1996, leg. M.G.V. Barbosa (BMNH) Paratypes: Brazil . 2 females from the same location as holotype, but collected on April and Aug. 1995, leg. M.G.V. Barbosa (BMNH); Peru. 1 female, Huanuco, Yuypichis, Panguana (74°56.8’W, 9°37’S), manioca field, 22.9.1975, leg. W. Hanagarth (UIC). Diagnosis. G. amazonensis is characterised by the deep punctation of the pronotum and the elytra. G. remelleti Orousset, 1985 has only free larger punctures in a longi- tudinal row close to the midline. The Chilean species can be distinguished from the Amazonian species by the prominent shoulders. Description. Length: 1.35 mm. Colouration: light brown, legs and antennae yellow. Head: 0.17 mm long, 0.22 mm wide; widest shortly in front of posterior margin; clypeus slightly cheek-like ex- tended; without distinctly narrower neck; without eyes; base of antennae to anterior edge; in dorsal aspect; ante- rior edge even; pair of setiferous punctures between base of antennae and laterally on posterior vertex; punctation weak; only few punctures larger; netlike micro sculpture weak; surface shiny. Antennae slightly longer than head and half of prono- tum combined; first antennomere thick; second oval and slightly narrower than first antennomere; third anten- nomere conical and slightly shorter than second; follow- ing antennomeres increasing in width except narrower sixth and eighth antennomeres; antennomere four to ten much wider than long; fourth antennomere twice as wide as long; tenth antennomere slightly wider than twice as wide as long. Pronotum: 0.23 mm long, 0.25 mm wide; widest near middle; sides evenly narrowed to anterior and posterior angles, but posteriad more strongly narrowed than ante- Bonn zoological Bulletin 64 (2): 109-116 ©ZFMK A new genus and new species of Neotropical Thoracophorini 115 riad; posterior angles obtuse; lateral margin continued to posterior edge; anterior edge not margined; punctation much deeper and coarser than on head; narrow midline impunctate; on average, interstices between punctures half as wide as diameter of punctures; in posterior half of mid- line with oval impression; netlike micro sculpture weak, but deeper than on head; surface moderately shiny. Elytra: 0.21 mm long, 0.28 mm wide; widest near pos- terior edge; narrowed to shortly rounded shoulders; punc- tation weaker and sparser than on pronotum; on average, interstices at least as wide as diameter of punctures; two pairs of large circular impressions close to suture; poste- rior impression larger than anterior impression; three to four setiferous punctures in lateral margin; netlike mi- crosculpture as deep as and as wide as on pronotum; sur- face moderately shiny. Abdomen as deeply, but more densely punctate than ely- tra; netlike microsculpture deeper than on pronotum and elytra; meshes distinctly wider than diameter of punctures; surface less shiny than on pronotum. Aedeagus with thick central lobe; central lobe with long slender and straight digit ending in acute curved apex; dig- it at ventral side with row of setae; paramera thick and sin- uate; several sensillae on ventral and dorsal side. Etymology. The specific name derived from the Brazil- ian state Amazonas, where the species was collected. Key to the species of Geomitopsis in the Neotropical re- gion 1 . Pronotum distinctly and densely punctate, elytra with round impressions close to suture (Fig. 5F) G. amazonensis n. sp. - Pronotum weakly and sparsely punctate, elytra with- out impressions 2 2. Shoulders of elytra not carinate, elytra posteriad not dilated (Fig. 5H) 4 - Shoulders of elytra carinate, elytra posteriad dilated (Fig. 5G) 3 3. Pronotum with large impressions on the disc G. campanae Saiz, 1973 - Pronotum with fine elongate impression in the pos- terior half of the midline G. chilensis Coiffait & Saiz, 1963 4. Aedeagus at ventral edge of central lobe with trian- gular prominence G. amrishi (Makhan, 2007) - Aedeagus at ventral edge of central lobe straight, without prominence 5 5. Central lobe of aedeagus at ventral edge straight, with short hook-like apex G rishwani (Makhan, 2007) - Central lobe of aedeagus at ventral edge evenly curved to acute apex G remilleti Orousset, 1985 DISCUSSION The new genus Geotrochopsis must be certainly placed to the tribe Thoracophorini, as the protibia has no inner emargination with comb such as in the tribe Osoriini. Re- garding the united gular sutures, it seems most closely re- lated to the genus Clavilispinus Bernhauer, 1926. In con- trast to the other blind genus, i.e. Geomitopsis, of the same tribe that has four tarsomeres, Geotrochopsis has five tar- someres. Furthermore, Geomitopsis was placed to the sub- tribe Glyptomina by Herman (2001), because gular sutures are separated. Irmler (2010) found that gular sutures can be separated or united even in one genus, which makes their constitution unsuitable for a generic differentiation. Nevertheless, the generic characters, e.g. tarsi composed of five tarsomeres and united gular sutures let suppose that Geotrochopsis is closely related to Clavilispinus, although the overall habitus is very different. Clavilispinus has well developed eyes, a dorsoventrally depressed body, and no pubescence, whereas Geotrochopsis is blind with cylin- drical body and dense pubescence. It might be also relat- ed to Allotrochus Fagel, 1955 regarding the overall habi- tus and the slightly elongate procoxae. But, Allotrochus has well developed eyes, no shortened elytra, and no pu- bescence. According to Herman (2001) the genus Geomitopsis is recorded from Europe, Africa, and South America. Nine species are loiown from the Mediterranean Europe includ- ing northern Africa and Near East, six from tropical Africa, and, together with the newly described species, six species from the Neotropical region. Whereas at present several blind genera are known in the tribe Osoriini, only this genus with blind species was so far known from the tribe Thoracophorini. The genus is characterised by the absence of eyes, tarsi composed of four tarsomeres, shortened ely- tra, a narrow sixth antennomere, and a characteristic struc- ture of the aedeagus. A similar combination of characters is found in the genus Arborilispinus Irmler, 2010. How- ever, Arborilispinus has eyes while eyes are absent in Ge- omitopsis. Acknowledgements. I thank the curators of the museums (L. Herman, R. Brooks, Z. Falin) as well as Jifi Janak for their steady support and for the loan of the specimens. Heartily thanks are also to A. Taghavian and J. Orousset from the Museum nation- al d’Histoire naturelle, Paris, for the loan of G. remilleti and the delivery of a paratype. I thank also D. Ahrens for the helpful comments to improve the manuscript. REFERENCES Makhan D (2008) Ashnaosorius gen. nov. from Suriname (Coleoptera: Staphylinidae: Osoriinae). Caloderma Suppl. 70: 1-8 Makhan D (2007) Two new species of Cubanotyphlus from Suri- name (Coleoptera: Staphylinidae: Leptotyphlinae). Caloder- ma 9: 5-8 Bonn zoological Bulletin 64 (2): 109-116 ©ZFMK 116 Ulrich Irmler Scheerpeltz O (1931) Zoologische Forschungsreise nach den Jo- nischen Inseln und dem Peleponnes. Sitzungsberichte Akade- mie der Wissenschaften, Wien 140; 359-460 Orousset J (1985) Deux Osoriinae anophthalmes de Guyane Frangaise (Coleoptera, Staphylinidae). Revue frangaise Ento- mologie (N.S.) 7; 60-66 CoifFait FI (1954) Contribution a la connaissance de la faune cav- emicole et endogee du Liban. (Mission H. Coiffait du Liban, 1951). IF Staphylinides nouveaux. Notes et Memoires sur le Moyen-Orient 5: 155-162 Coiffait H, Saiz F (1965) Une espece du genre Geomitopsis (Col. Staphylinidae). Revue Ecologie Biologie du Sol 4: 577-578 Saiz F (1973) Un nouveau Geomitopsis du Chile (Col. Staphy- linidae). Nouveau Revue entomologique 1; 19-21 Herman E (2001) Catalogue of the Staphylinidae (Insecta; Coleoptera). 1758 to the end of the second millennium. II. Oxyteline group. Bulletin of the American Museum of Natu- ral History 265: 1067-1806 Irmler U (2010) A new genus of Osoriinae in the Neotropical region with a cladistic analysis of the tribe Thoracophorini (In- secta: Coleoptera: Staphylinidae). Arthropod Systematics & Phylogeny 68: 229-237 Bernhauer M (1926) 32. Beitrag zur Staphylinidenfauna Siid- amerikas insbesondere Brasiliens. Memoire della Societa En- tomologica Italiana 5; 152-169 Fagel G 0955) Osoriinae. Pp. 1-133 in: de Witte, G.F. (ed.) Ex- ploration du Parc National due FUpemba. Institut du Parces nationaux du Congo Beige 39 Bonn zoological Bulletin 64 (2): 109-116 ©ZFMK Bonn zoological Bulletin 64 (2): 117-138 March 2016 A critical review of Hoser’s writings on Draconinae, Amphibolurinae, Laudakia and Uromastycinae (Squamata: Agamidae) Wolfgang Denzer* ^ Ulrich Manthey^ Philipp WagneU^ & Wolfgang Bohme^ ^ Society for Southeast Asian Herpetology, Rubensstr. 90, D-12157 Berlin, Germany; E-mail: wolfdenoxford@yahoo.co.uk ^ Society for Southeast Asian Herpetology, Kindelbergweg 15, D-12249 Berlin, Germany; E-mail: manthey. sseah@t-online. de ^ Zoologische Staatssammlung Munchen, Munchhausenstr. 21, D-81247 Munchen, Germany; E-mail: philipp. wagner.zfmk@uni-bonn. de Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA; ^ Zoologisches Forschungsmuseum Alexander Koenig, Leibniz-lnstitut fur Biodiversitat der Here, Adenauerallee 160, D-53113 Bonn, Germany; E-mail: w.boehme@zfmk.de ^ Corresponding author Abstract. We analyzed four papers on agamid lizards by self-proclaimed Australian herpetologist Raymond Hoser with respect to the presentation of diagnostic characters as well as their taxonomic and nomenclatural merits. In most cases the taxonomic concepts were lifted from earlier phylogenetic publications and the diagnoses were copied from other au- thors. Copied text in Hoser’s diagnostic section within the analyzed papers amounts to a staggering 83% for Draconinae, 82% for Amphibolurinae, 77% for Laudakia and 78% for Uromastycinae, respectively. We found a number of plagia- rized paragraphs, sometimes half a page long. Hoser hardly ever makes any effort to attribute statements to the original author and in some cases he even omitted to cite the relevant source. With respect to nomenclature, we found that Hoser proposed names that were preoccupied or unavailable, that a nomen oblitum was resurrected incorrectly, nomina nuda were produced, a type locality was restricted incorrectly and a questionable holotype was designated for a new species. With respect to taxonomy, we found examples of wrong diagnoses, falsely attributed species, omission of taxa and a lack of understanding or misinterpretation of previously published taxonomic studies on agamid lizards. Furthermore rele- vant literature on taxonomy and nomenclature has been overlooked or disregarded. Key words. Plagiarism, IZCN rules, nomina nuda, questionable type specimen designation, ambiguous diagnoses INTRODUCTION For the past few years now the Australasian Journal of Herpetology (hereafter AJH) has been produced in print and as an online journal where pdfs can be downloaded. At the time of writing, 29 issues of the AJH have been produced. The editor of and sole contributor to the jour- nal appears to be Raymond Hoser who mainly writes about reptile classification. These articles are an area of contro- versy and most herpetologists as well as herpetological journals and societies worldwide have recorded their ob- jection to Hoser’s works (see Items for Action & Aclmowl- edgments in Kaiser 2013); the scientific community cur- rently appears almost unanimous in their approach not to use Hoser’s nomenclature. Albeit that the majority of herpetologists appears to be in agreement on the suggested suppression of names pro- posed by Hoser, it has to be noted that this action may not be in agreement with The International Code of Zoolog- ical Nomenclature (ICZN, 1999 & 2012; hereafter “the Received: 11.08.2015 Accepted: 22.03.2016 Code"'), a set of regulation every zoologist is obligated to follow and should wish to uphold. The Code is served by the International Commission on Zoological Nomencla- ture (hereafter, ICZN), which adjudicates instances where taxon names may lead to confusion, are improperly pre- sented or formed, or where published works threaten the stability of the nomenclature in a given discipline. The service of the ICZN includes a recently developed, for- mal taxon name registration service in the form of Zoobank (accessible at zoobank.org), where authors of taxon names may formally establish a claim to their names or other nomenclatural acts. Hoser registers all names pro- posed by him with Zoobank and as a consequence the names are available in the sense of the Code. However, it must be noted that the Zoobank website does not have any provision to prevent the registration of invalid nomen- clatural acts, thus anyone can register and contribute pre- sumed valid scientific names. In its current version. Corresponding editor: F. Herder 118 Wolfgang Denzer et al. Zoobank can only be considered as provisional until there are rules implemented that prevent misuse of this data- bank. The Code has no provisions for the quality of publica- tion in which taxonomic and nomenclatural acts are pro- posed. In particular, there is no need for a journal to have an editorial board or have a peer review process in place to validate a proposed name. As has been noted, “the qual- ity of taxonomic descriptions does not make a name un- available there being no requirement as such in the Code..."" (Thomson 2014), i.e. for the ICZN nomencla- ture and taxonomy are not dependent upon each other. A proposed taxonomy may be inconsistent, ambiguous or even false and every herpetologist can choose to follow it or not, but a proposed taxon name, if produced in ac- cordance with the Code, becomes available immediately. There exist only a few prerequisites for a journal to com- ply with the Code in order to validate and make available a proposed name. One such prerequisite (ICZN, Article 8) is that the journal is widely available (for example in public libraries) “providing a public and permanent sci- entific record” and “numerous identical and durable copies” have to be assured. Typically 25 copies (Recom- mendation 8b) constitute a sufficiently available edition. In order to prove that sufficient copies have been printed Hoser typically publishes a tax invoice in each issue of AJH stating that 50 copies were printed. Distribution is, however, not proven, but presumably at least some copies are sent to libraries (all issues of the journal can be found in the National Library of Australia online catalogue) and distributed among subscribers to the journal. Additional- ly, all issues or individual articles within a given issue are presented online as downloadable pdfs a month after the print version has been in circulation. Every nomenclatur- al act is registered with Zoobank and hence the proposed names may be considered published in accordance with the Code and therefore available for the purposes of nomenclature. Editorial boards and high profile referees (reviewers) of manuscripts are usually a measure for the quality of a jour- nal and their names may even be published periodically (e.g. Journal of Herpetology). The AJHdoQS not have an editorial board to oversee standards of publication or for undisclosed reasons has decided not to present that infor- mation in any issues of AJH. However, according to Hoser (2012: 41) manuscripts submitted to the journal are ref- ereed by four independent reviewers. This extensive peer review process should assure that all taxonomic and nomenclatural decisions presented “[stand] up to the most robust of scrutiny” (Hoser 2012: 41). Additionally this lev- el of peer review should provide an assurance that the ar- ticle adheres to commonly accepted editorial standards, including ethical considerations such as avoidance of pla- giarism or the inclusion of derogatory comments. Plagiarism is generally defined as passing off ideas or text from other publications as one’s own, whether or not the source is cited (for definitions see plagiarism.org). Copying text into one’s own work without citing its source is the most flagrant form of plagiarism and in many coun- tries is a violation of intellectual property rights and ille- gal. Even copying a substantial part of a previous publi- cation and citing the source is still a form of plagiarism, if the copied text is not produced within quotation marks or other means to make the reader aware that the original research or text is not the work of the current author. Sim- ilarly, minor modification of the original text such as re- arrangement of phrases or the substitution of a few words is still plagiarism, when the original author is not attrib- uted in an appropriate manner. Derogatory criticism of other authors in any scientific publication must be avoided. Providing counterarguments relating to scientific opinions of a certain author or a group of authors is a well-established way in science to encour- age discussion about the matter in question. However, per- sonal attacks or defamations must be avoided by all means and are not a part of a scientific (or other) publication. In the following discussion we will analyze four of Hoser’s (Hoser 2012a, 2013, 2014b & 2014c) publications on agamid lizards and discuss our findings in taxonomic and nomenclatural terms. MATERIALS & METHODS The papers were downloaded from the AJH website. Hoser’s texts were analyzed with respect to their taxonom- ic and nomenclatural decisions as well as to generally ac- cepted editorial standards of scientific publications. Pre- vious publications by other authors containing diagnos- tic characters and descriptions were compared to the di- agnoses used by Hoser. Any copied or plagiarized text was marked and attributed to the original source including page number. Hoser’s diagnoses do not follow Einnean tele- graphic style and frequently contain long introductory sen- tences that do not further the knowledge about a taxon. We, therefore, accounted for any copied or plagiarized text identified in Hoser’s diagnoses in two different ways: 1) as a percentage of the whole diagnosis including introduc- tory sentences and 2) as a percentage of the presented text comprising diagnostic characters only. This was done by accounting for lines of overall text vs. lines of copied text in a way that favoured any originality in Hoser’s text, i.e. a line, even if only half printed, was typically counted as full, while in the case of copied text two half lines were counted as one. Total lines in the publication about Am- phibolurinae (Hoser 2013) were counted, those of the oth- er publications discussed here were estimated as follows: typically each page in the A J// contained about 140 lines (70 lines per column). Abstracts and titles were printed in Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK Critical review of Hoser’s publications on agamid lizards 119 full lines and the actual number of lines was therefore dou- bled as if they had been in two columns. In the case of the other papers the overall sum of lines was not count- ed but calculated by assuming that each column contains 70 lines. One of the sources referenced by Hoser (2013) is Cog- ger (2000). Here we present the results in comparison to Cogger (1983) in order to show that nearly all of the di- agnostic characters used for the classification of amphi- bolurine lizards are considerably older than claimed. Some diagnostic characters could not be accounted for by com- parison to earlier publications. Where the source was un- clear an internet search was performed and if identified (e.g. Wikipedia, Reptile Database etc.) parts were marked accordingly. Obviously we do not know precisely which sources were actually used by Hoser (original description, review works, catalogues, web pages etc.) and therefore we relate identified text passages to the publication where we looked for and found identical phrases. As we cannot reproduce every single character or paragraph for direct comparison the respective pages where sets of characters or a full description can be found are given together with the number of copied lines and the respective source. At the end of each section we give a summary of our find- ings with informations on Hoser’s taxonomical approach and sources used. DISCLAIMER As a general rule Hoser’s new taxon names are not used in this paper and the respective taxon named by Hoser will be mentioned as “new tribe / genus to accommodate / con- tain the following XY” or by a similar phrase where the placeholders are substituted by currently accepted names. This is done to prevent accidental validation of Hoser’s names, which subsequently could become available un- der the rules of the Code. If, by accident, a new taxon name proposed by Hoser is used herein that paragraph shall be treated as not published and the name shall be con- sidered as not available for the purposes of nomenclature. This disclaimer is in compliance with Article 8.2 of the Code. RESULTS & DISCUSSION A) Hoser (2014b) on Draconinae As printed in the header of the paper, the Draconinae man- uscript was received by the journal on 10 November 2013, accepted on 1 June 2014 and published on 1 July 2014. According to the tax invoice. Issue 22 of the AJH, which includes the Draconinae paper, appears to have been planned before October 2013, which is the date of the in- voice (Hoser 2013: 36, Hoser 2014a: 5; invoice date 3 Oc- tober 2013, several weeks before the publisher initially re- ceived the manuscript). This could indicate that Hoser pays in advance for the printing of issues, which would imply that manuscripts may already be in hand, or that some of the publication dates are otherwise manipulated. The paper contains the following sections or headings: Title, Abstract (including Keywords), Introduction, Unlaw- ful Theft of Material and Data, and Notes on Taxa Named Herein, followed by the actual taxonomic and nomenclat- ural part, a Conflict of Interest section, and a References Cited section. The publication additionally contains a table depicting the proposed nomenclature. The introduction to the paper is mainly concerned with the phylogenetic and morphological data presented by ear- lier authors, which serve as the basis for Hoser’s taxonom- ic and nomenclatural decisions. As in most of his recent papers, Hoser includes personal criticism of recent and past herpetologists. Similarly, Hoser directly insults sev- eral herpetologists in his Unlawful Theft of Material and Data section of the paper. In this part we are also made to believe that most of his research files had been confis- cated and that his ideas were repeatedly used by recent authors in order to rename taxa and produce junior syn- onyms. Overall, in this paper Hoser describes one new species, proposes eight new genera, resurrects three names for sub- genera, and erects 22 subgenera, ten new tribes and six subtribes. His diagnosis of the genus Lyriocephalus Mer- rem, 1820 may serve as an example how he defines a genus and how we analyzed his statements. The follow- ing is a true copy from Hoser (2014a: 38): “Lyriocephalus Merrem, 1 820 is defined by the follow- ing suite of characters: Mouth large; teeth erect in both jaws. Incisors small and conical. No praeanal or femoral pores (as opposed to the callous pore-like swelling of the preanal scales of the males in the genera Agama Daudin, 1802, UromastixMQrrQm, \^20 andXenagamaBoulQngQY, 1895); tympanum hidden. Five toes. A dorsal crest; a V- shaped gular fold; a bony supraorbital arch. Body com- pressed, covered with small scales intermixed with en- larged ones. A nuchal and a dorsal crest. A gular sac and a V-shaped gular fold. Adult with a globular hump on the nose. Pre and post-orbital bones forming an arch limiting a supraorbital fossa.” The first set of characters “Mouth large . . . arch” is a copy from Boulenger’s synopsis leading to Lyriocephalus (Boulenger, 1885: 251-252). The part in brackets “callous . . . genera” is taken from a footnote in Boulenger (1885: 251), where it only refers to Agama and Aporoscelis [= Xenagama]. The part containing Uromastix [sic] andXe- nagama could not be identified, but is presumably taken from another comparatively old publication as the genus name Uromastyx is written in its historically used form. The second set of characters “Body ... fossa” mirrors Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK 120 Wolfgang Denzer et al. Boulenger’s (1885: 281) diagnosis of the genus. It is quite obvious that copying has been done without giving it much further consideration. The V-shaped gular fold appears twice as does the dorsal crest. The characters “supraor- bital arch” and “supraorbital fossa” are repeated without comment; when a supraorbital arch is formed, this leads to a supraorbital fossa between the arch and the dorsal out- er ridge of the eye socket. Hoser cites Moody (1980) in his bibliography. Had he looked at this publication he would have found that as a matter of fact the supraorbital arch in Lyriocephalus is formed by prefrontal and postor- bital and not as claimed by Hoser (2014a: 38) by “pre and post-orbital bones”. Most taxonomic concepts proposed by Hoser have been published by earlier authors (and cannot be repeated here in full for comparison) but without taking the step of as- signing genus names to species groups (e.g., Gono- cephalus Kaup, 1825; Draco Linnaeus, 1758; Japalura Gray, 1853) or to species where only insufficient materi- al and/or data exist. In the following we will first provide evidence that the taxonomic scheme proposed by Hoser is either based on previously published concepts or con- stitutes mere naming of more or less supported nodes in phylogenetic publications concerned with Draconinae. In the second part we will have a closer look at the diagnoses of genera and compare those to previously published ma- terial. We will discuss each group in the same sequence as published by Hoser. In our analysis below we will not discuss all of Hoser’s diagnoses in such great detail as the one of Lyriocephalus and only point out inconsistencies in taxonomy and nomenclature where we feel it should be done for clarity. The first genus Hoser deals with is Gonocephalus which he proposes to divide into five subgenera along with the erection of two new genera. His subgeneric classification follows the species group assignment proposed by Man- they & Denzer (1991) and Denzer & Manthey (2009, part). Denzer & Manthey (l.c.) combined the Philippine species with their bornensislbellii species group, which they had considered a separate species group in the earlier publi- cation, based on morphological similarities. Hoser elevates two species to genus rank, namely G. robinsonii (Boulenger, 1908) and G. mjobergi Smith, 1925. This had already been suggested by Manthey (2010) where G robinsonii was treated as (Gonocephalus incertae sedis) robinsonii and by Denzer & Manthey (l.c.) where it was suggested that G. mjobergi should be referred to as Genus A within a Gonocephalus s. 1. complex. Owing to insuf- ficient material (only a single female specimen has ever been collected) Denzer & Manthey (l.c.) abstained from proposing a genus name for G mjobergi until more ma- terial will become available. They further stated that one autapomorphic character in particular (longitudinal gular folds) constituted a synapomorphy for the genus group G mjobergi, Mantheyus Armyowdi & Stuart, 2001 and Ptyc- tolaemus Peters, 1 864. With respect to G robinsonii Hoser states in his introductory part to the genus Gonocephalus that “no one has bothered to assign the taxon Gono- cephalus robinsonii ... to a genus of its own”. As will be discussed below Hoser has taken on this task but fails to deliver as not a single of his characters is of any value to diagnose his newly proposed genus (i.e., differentiate from other genera or species groups). Initially Hoser characterizes the genus Gonocephalus. His diagnosis comprises seven lines and is copied from Boulenger (1885: 282, 3.5 lines) and Denzer & Manthey (2009: 255-256, 3.5 lines). His subgenus to accommodate the chamaeleontinus group as defined by Manthey & Den- zer (1991) is characterized by a single character copied from Boulenger (1885) and separated from other proposed subgenera by comparison in a way that their full diagnoses are repeated. Hoser considers the chamaeleontinus group as the nominate form. Next Hoser proposes a subgenus to accommodate G grandis (Gray, 1845). For this species he resurrects an available name proposed by Gray (1845). The diagnostic character section amounts to ap- proximately 25 lines, all of which are a copy of Boulenger’s (1885: 298) description of the species. This is followed by proposing a new subgenus for the Philip- pine species group by using three characters (two lines) taken from Boulenger (1885). The next new subgenus comprises the bornensis group. The diagnostic characters are mainly taken from Boulenger (1885) but rearranged and slightly modified without copying directly. The name he gives the subgenus is different from the name he uses in the keywords to the paper, the latter of which therefore becomes a nomen nudum. The last new subgenus proposed contains the Sumatran megalepis species group and is characterized initially by two lines copied from Boulenger’s (1885) synopsis [key] to the genus followed by Boulenger’s (1885: 291) full description of G tuber- culatus (= G megalepis). This last part comprises 24 copied lines and ends with citing Boulenger (1885). How- ever, Hoser does not make clear that the whole descrip- tion is copied by, for example, using quotation marks. Gonocephalus robinsonii is removed from its synonymy with the genus and a new genus is proposed. This new genus is diagnosed by three characters: karyotype, a “greatly enlarged gular fold” and “a distinctive white low- er jaw”. The karyotype section is a copy from Diong et al. (2000: 74, 6 lines); the other two characters are sup- posedly based on Hoser’s own research. We would like to note that the karyotype can even vary within a species (e.g., see Ota, 1988 for data on Japalura swinhonis Guen- ther, 1 864), the enlarged gular fold is a false character as G robinsonii is the only Gonocephalus species without a gular fold, if one considers G mjobergi as not congener- ic and the colour of the lower jaw constitutes a variable character in G robinsonii which is dependant on age (see photographs in Manthey 2010). Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK Critical review of Hoser’s publications on agamid lizards 121 Gonocephalus mjobergi is accommodated in a new genus. This is done by copying the full description from Denzer & Manthey (2009, 40 lines) including the above- mentioned paragraph about the autapomorphy of longi- tudinal gular folds. Despite using the complete character set including that a large gular sac “partially conceals the Gonocephalus-typQ typical gular fold” Hoser earlier claims that G. robinsonii and G mjobergi have an “en- larged gular fold.” Additionally this quote shows that Hoser apparently intended to amend the original statement but ended up doubling an adjective. The original phrase reads: “partially conceals the Gonocephalus gular fold” (Manthey & Denzer 2009: 257). Hoser often uses brackets for the author of a taxon and the year of description where he seems to interpret the Code in his own way (e.g. he uses brackets for Gono- cephalus robinsonii, Boulenger, 1908, G beyschlagi Boettger, 1892, G doriae Peters, 1871). The use of brack- ets for the author/year of a taxon is determined by the ICZN rules (Article 51.3 [Use of parentheses], see ICZN Code for an example). The Code prescribes brackets if the allocation of a species changes with respect to a genus. This is not the case here. Boulenger and the other authors decided that the correct spelling should be Gonyocephalus (an emendation introduced by Wagler [1830]) albeit that Kaup originally used Gonocephalus and later amended it to Goniocephalus, but the latter emendation and Gony- ocephalus are not available under ICZN rules as the orig- inal name has to be preserved. The genus Japalura Gray, 1853 is broken up into three genera, two of them divided additionally into two subgen- era each. Japalura has for a long time been a matter of taxonomic changes and only in recent years are we be- ginning to understand their phylogenetic relationships. A division into three genera can be derived from molecular phylogenetic analyses, where results indicate that the clades containing J. variegata Gray, 1 853 / J. tricarina- ta (Blyth, 1853), J. polygonata (Hallowell, 1861), and J. splendida Barbour & Dunn, 1919 / J. flaviceps Barbour & Dunn, 1919 are only very remotely related (e.g., Py- ron et al. 2013). Stuart-Fox & Owens (2003) considered Japalura as comprised of “two widely divergent goups,” named in their analysis as Japalura India / J. variegata- group and Japalura SE Asia / J. splendida-growp (SE for Southeast). They even mention that they consider both as separate genera. Mahony (2009: 55) refers to the latter species group as “eastern elade”. In an earlier publication by Macey et al. (2000) they are referred to as Himalayan and East Asian clades, respectively. Kastle & Schleich (1998) proposed that the species of the Western clade with a visible tympanum should be regarded as a separate genus, for which the name Oriotiaris Gunther, 1 864 was available. Hoser mostly follows these previously published results to propose his taxonomic scheme. Firstly he deals with Japalura species of the nominate genus. Here he seems to accept Mahony’s view (2009) that Japalura and Oriotiaris are congeneric. Japalura varie- gata (type species of Japalura) and J. tricarinata (type species of Oriotiaris) are phylogenetically sufficiently close (Pyron et al, l.c., papers cited in Mahony, l.c.) that Mahony (2009) already suggested to synonymize both genera and treat Oriotiaris (resurrected by Kastle & Schle- ich (1998)) as a Junior synonym of Japalura. Hoser treats both as subgenera of Japalura. The nominate genus Japalura is diagnosed in seven lines which are copied from Boulenger (1885: 307) and Mahony (2010: 4, definition of Japalura s.l.) with approx- imately half of the text from each author. The genus is fur- ther divided into a nominate subgenus and by resurrect- ing an available name for the second subgenus. Hoser first defines Oriotiaris. His diagnostic characters for this sub- genus are copied from Gunther (1864: 1 50, five lines) and Mahony (2009: 56, five lines). No other characters are giv- en. In the case of one character taken from Mahony (2009) Hoser even copies a typographic error, “. . .possession of a small gular pouch in the later” [sic!]. The subgenus Japalura is diagnosed as follows: “The diagnosis for the nominate subgenus Japalura is simply a reversal of the diagnosis for Oriotiaris.'' Hoser distin- guishes Japalura from his subgenus Oriotiaris as follows: ''Oriotiaris is further separated from the nominate sub- genus Japalura by the absence (vs. presence) of dorsal chevrons and presence (vs. absence) of a coloured gular region, concealed tympanum, large crest spines in males and erectile nuchal crest (roach), in members of Japalu- ra." Japalura tricarinata is highly variable and eapable of changing colour. There exist photographs of complete- ly green individuals without any chevron pattern (see for example Manthey 2010: 98, Fig RA02806-4). On the oth- er hand, J. planidorsata Jerdon, 1870 does not have an erectile nuchal crest nor does J. sagittifera Smith, 1 940 both of which are placed by Hoser in the nominate sub- genus. For Japalura polygonata Hoser resurrects its original name Diploderma polygonatum Hallowell, 1861. Phylo- genetic studies showed that J. polygonata is only remote- ly related to other Japalura but seems to be the sister tax- on of Gonocephalus robinsonii (Pyron et al. 2013). Diplo- derma polygonatum was already suggested by Mahony (2009: 55) in case the eastern clade (see below) should turn out to be monophyletic. The genus is diagnosed with four lines, all of which are copied from Boulenger (1885: 307). Having already dealt with the variegata group, Hoser proceeds to define a new genus for the eastern species group, which he splits into two subgenera. The new genus to accommodate all East Asian species is defined in the space of approximately nine lines, seven of which are directly copied from Boulenger (1885: 307-308). One Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK 122 Wolfgang Denzer et al. character is the negation of a Boulenger character and the rest are slightly amended but not identically copied from Mahony (2009). No new characters are introduced by Hoser. This genus is split into two subgenera on the ba- sis of Boulenger’s synopsis (1885:308) the differences be- ing the length of the tibia and presence or absence of a longitudinal fold. For J. swinhonis, Hoser claims that the “tibia is as long as the skull”. Already Stejneger (1907: 1 83) pointed out that in two old males he had studied “the tibia is decidedly shorter than the skull”. With respect to the second character we would like to note that J. chapaen- sis Bourret, 1937, J. fasciata Mertens, 1926, J. grahami (Stejneger, 1924) and J. micangshanensis Song, 1987 do not have a longitudinal fold as claimed by Hoser. They would therefore have to be transferred to his first sub- genus. The genus Calotes Daudin, 1 802 is divided into three genera, of which two are further divided into subgenera (the nominate genus into five and a newly proposed genus into four subgenera). The basis for this taxonomic scheme appears to be the result of the extensive molecu- lar biological studies presented by Zug et al. (2006) and Pyron et al. (2013). The proposed scheme clearly reflects the nodes in previously published phylogenetic trees. A division into three groups was already proposed by Smith (1935) who differentiated between a C. versicolor group, a C. liocephalus group and a group comprising C. ronxi Dumeril & Bibron, 1837 and C. ellioti Gunther, 1864. The first and last of Smith’s groups are elevated to genus lev- el by Hoser, the liocephalus group is considered by Hoser as a subgenus. Initially the genus Calotes is diagnosed by a copy of Boulenger’s diagnosis (1885:314, four lines) and subse- quently compared to his newly erected genera (see below). The first subgenus described within Calotes serves to ac- commodate C. calotes (Linnaeus, 1758) and C. htunwini Zug & Vindum, 2006. Their close relationship was dis- covered in phylogenetic studies despite the fact that their distribution is rather disjunct. The nominate subgenus is defined by Hoser by initially repeating Boulenger’s key (1885: 315-316, 3.5 lines) leading to C. ophiomachus (= C. calotes) and subsequently by a complete copy of Boulenger’s description of the species (Boulenger 1885:327, approximately 18 lines). By stating that all of these characters define the subgenus Hoser renders his di- agnosis false. C. htunwini does not have a nuchal crest where the height “equals or exceeds the diameter of the orbif ’ nor does it have a “dorso-nuchal crest composed of closely set lanceolate spines” nor is this species green above. Additionally we like to note that already Boulenger’s description contains a mistake in stating that in C. calotes a “gular sac is not developed”. This has been copied by Hoser; however, male C. calotes actually have a reasonably well developed gular sac during the breed- ing season as have C htunwini but to a lesser extent. Next Hoser proposes a new subgenus containing species allied to Calotes versicolor (Daudin, 1802) by copying 1.5 lines from Boulenger’s synopsis (Boulenger 1885: 314-315) followed by an entirely copied descrip- tion of C versicolor from Boulenger (1885: 312, 20 lines). Here also Hoser repeats Boulenger’s statement that in C. versicolor the “gular pouch [is] not developed” which is not true for male specimens during the breeding season (Smith 1935; numerous photographs on the internet). Hoser does not present any new characters for the C. ver- sicolor group. Subsequently Hoser erects a new subgenus containing two closely related species from the Western Ghats, namely C. nemoricola Jerdon, 1853 and C. gran- disquamis Gunther, 1875. To diagnose the genus he ini- tially copies four lines from Boulenger’s synopsis (1885: 315) leading to these species followed by the reproduc- tion of Boulenger’s description (1885: 326) of C. nemori- cola (approximately 20 lines). No additional or new char- acters are presented by Hoser. Species allied to Calotes liolepis Boulenger, 1885 (C. nigrilabris Peters, 1860 and C. desilvai Bahir & Maduwage, 2005) are the content of a subgenus that is ini- tially defined by repeating in full Hallermann’s key (2000: 161-162) leading to C. nigrilabris and C. liolepis, respec- tively (3.5 lines each), followed by a copy of Boulenger’s descriptions (1885: 327-328) of C. nigrilabris (approxi- mately 22 lines) and C. liolepis (approximately 15 lines). No other characters are presented in the diagnosis. Species related to Calotes liocephalus Gunther, 1 872 are placed by Hoser into a new subgenus which again is de- fined by the characters given in Hallermann’s key (2000) here for C. liocephalus and C. ceylonensis Muller, 1887 (3.5 lines each) followed by the respective descriptions copied from Boulenger (1885: 329) for C. liocephalus (18 lines) and Boulenger (1890: 139-140) for C. ceylonensis (13 lines) without presenting any further characters. The last subgenus within Calotes proposed by Hoser is monotypic and contains only C. aurantolabium Krishnan, 2008. Diagnostic characters are given in the space of 13 lines all of which are a copy of Krishnan (2008). After having dealt with the species he considers Calotes sensu stricto. Hoser proceeds to erect a new genus to accommodate species related to C. mystaceus Dumeril & Bibron, 1837. This genus is further divided into four subgenera. The definition of the genus comprises approx- imately 13 lines, which are a copy from Boulenger (1885: 315) or may partially have been taken from Hallermann (2000: 162). Initially Hoser gives a short diagnosis for the genus (2.5 lines) followed by the sentence: “In addition to this, each of the relevant subgenera are further diag- nosed and separated from the other genera by one or oth- er of: A/ [diagnosis subgenus A] or B/ [diagnosis subgenus B] or C/ [diagnosis subgenus C]”. This is followed by sep- arating the genus from Calotes and another genus contain- ing C. rouxii Dumeril & Bibron, 1 837 and C. ellioti Giin- Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK Critical review of Hoser’s publications on agamid lizards 123 ther, 1864. The presentation of his diagnoses for the sub- genera here is peculiar if not unique: the diagnostic char- acters including comparisons presented for the nominate subgenus and two other subgenera are absolutely identi- cal to that of the genus! The last subgenus is monotypic and erected for Calotes nigriplicatus Hallermann, 2000. Here he repeats the full description as given by Hallermann (2000: 156, 158, ap- proximately 30 lines) only adapted in places where a com- parison is made to one of his newly erected subgenera (i.e. the name Calotes is replaced by Hoser’s new name). This is followed by repeating again his diagnostic characters for the already defined subgenera and genera. In the space of two pages he uses the same 22 lines five times. In all of this Hoser does not present a single new character. Next he defines a new genus to accommodate Calotes rouxii and C. ellioti. The diagnosis comprises two lines and is copied from Hallermann’s key (2000: 162). The last genus Hoser proposes is again monotypic and only contains what he calls Calotes andamanensis, cur- rently considered as Pseudocalotes andamanensis (Boulenger, 1891). While Harikrishnan & Vasudevan (2013: 11) state: “.. .these differences are not sufficiently pronounced to justify the recognition of a new genus. In the absence of a molecular phylogeny and based on ex- ternal morphology alone, it is most appropriate to consid- er this species as a member of Pseudocalotes...'' Hoser opposes this by writing “is also sufficiently divergent to warrant being placed in a separate genus”. Hoser’s initial diagnosis is a complete copy (31 lines) from Krishnan’s description (2008: 533) of the species, only substituted with Hoser’s nomenclature in places where Krishnan made comparisons with Calotes. This is followed by the descrip- tion of Pseudocalotes andamanensis (14 lines) given by Harikrishnan & Vasudevan (2013: 11) and subsequently by yet another short description of this species including comparisons with Calotes Daudin, 1802, Bronchocela Kaup, 1827, Complicitus Manthey in Manthey and Grossmann, 1997, Salea Gray, 1845, and Dendragama Doria, 1888 (17 lines) as produced on the Reptile Data- base website (original publication not identified). We note that also the first two descriptions are available on the Rep- tile Database website. Hence Hoser could have copied the whole diagnosis from there without even consulting the original publications. This assumption is viable as Harikr- ishnan & Vasudevan (l.c.) are cited in an identical place to that on the website and Krishan’s description stays with- out a citation as this is also the case on the website. Al- together he “describes” the species three times in 65 lines of which 62 lines are copied from other sources and the remaining lines are introductory sentences. The genus Ceratophora Gray, 1835 is divided into three genera including two subgenera reflecting the molecular and morphological (rostral horn appendage) phylogeny of Schulte et al. (2002). The nominate genus contains the species related to C. stoddartii Gray, 1 834 which is divid- ed subsequently into two subgenera. Hoser’s description of the nominate genus is presented in 6.5 lines all of which are taken from Boulenger (1885: 277) with only minor changes. This is followed by a separation from his other proposed subgenus and the other two proposed genera (12 lines). The complete text to describe the diagnostic char- acter is copied from Boulenger (1885: 277) and Pethiyago- da & Manamendra-Arachchi (1998: 1,4). The definition of his subgenus to accommodate C tennentii Gunther, 1861 comprises approximately four lines all of which are taken from Boulenger’s synopsis (1885: 277). The nom- inate subgenus is defined by four lines again from Boulenger (1885: 277). Next Hoser erects a new genus for Ceratophora aspera Gunther, 1 864, which is initially defined by two lines from Boulenger (1885: 277) followed by characters taken from Pethiyagoda & Manamendra-Arachchi (1998: 44, 46, six lines, all copied) to separate it from the other proposed genera by Hoser. Even the distributional data are copied verbatim from Pethiyagoda & Manamendra-Arachchi (1998: 44). The last genus Hoser proposes for this group of lizards only contains Ceratophora karu Pethiyagoda & Mana- mendra-Arachchi, 1998. This is presented including comparisons within approximately eight lines, all of which are a copy from Pethiyagoda & Manamendra-Arachchi (1998: 44) and can partially be found in an identical way on the Reptile Database website. Next Hoser deals with the lizards of the genus Bron- chocela Kaup, 1827. He initially gives an introduction where he seems to restrict the type locality of B. cristatella (Kuhl, 1820) and to resurrect B. moluccana (Lesson, 1830) (see discussion below). The genus is di- vided into two subgenera the first of which contains B.ju- bata (Dumeril & Bibron, 1 837) and B. orlovi Hallermann, 2004. The first three lines of the diagnosis are taken from Boulenger (1885: 314 all copied) and a full description (approximately 20 lines) of B. jubata is presented by a copy of Hallermann’s description (2005: 171-172). Two more lines of characters concerning the scales at the base of the dorsal crest could have been taken from de Rooij (1915: 123). One character cannot be retraced to earlier publications and presumably comes from Hoser’s re- search: “The dorsal crest gives the appearance as if it is composed of tiny hairs as opposed to scales (as seen in Bronchocela)" [sensu Hoser]. We note that adult males of B. jubata have one of the most developed dorsal crests among Bronchocela, consisting of lanceolate scales. The only new species described by Hoser within the Draconinae paper is a member of Bronchocela Kaup, 1 827 and refers to material collected on Halmahera Island, Maluku Province, Indonesia. His description of this species is purely based on colouration and an elongated scale between the nasal and the rostral. We note that most Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK 124 Wolfgang Denzer et al. - if not all - Bronchocela species are capable of extreme colour changes. A typically brightly green coloured B. cristatella (Kuhl, 1820) may become completely black when disturbed or during copulation (WD pers. obs.). Hoser’s choice of holotype (USNM 237431) is - to put it mildly - slightly confusing. The specimen he chose ac- tually has a bifurcated tail something that should have been noted in the diagnosis (see collections. si.edu/search/re- sults.htm?q=record_ID:nmnhvz_6091296). We further note that the gender of Bronchocela is female but Hoser creates a species name with a masculine ending. The de- scription of his nQ^N Bronchocela species contains copied sections from Boulenger (1885: 314, 316-317, approxi- mately 26 lines) for B. cristatella. In his comparison of the new species to other species of the genus Bronchocela the author also often refers to B. moluccana (Lesson, 1 830) which is currently consid- ered a synonym of B. cristatella (Kuhl, 1820). Interest- ingly he does not include B. moluccana in his species list (table at the end of his taxonomic section) although it is stated in his introduction to the genus that he regards B. moluccana “as being a separate species”. We note that the original name given would be Agama moluccana Lesson, 1 830 and the combination B. moluccana was only used by Peters (1867 as Bronchocele), Stoliczka (1870) and Pe- ters & Doria (1878), all of which were in later publica- tions considered to be B. cristatella. Theobald (1876) used the name B. moluccana in his Reptiles of British India for a specimen from the Nicobars as a synonym of Pseudocalotes archiducissae Fitzinger, 1 860, which again turns out to be a synonym of B. cristatella. Bronchocela moluccana constitutes a nomen oblitum and resurrection should have been made clear with reference to the type species and holotype. Furthermore Hoser refers several times to Java as the type locality for B. cristatella. In his original description Kuhl (1820) never mentions a type locality and ever since it has been unknown and never been restricted by any au- thor (see for example Diong & Lim, 1998). One could ar- gue that Hoser’s statement “West Java (herein treated as terra typica)” is meant to be the newly defined type lo- cality. This is an unfortunate choice under current condi- tions, as the actual phylogenetic status of the Javanese pop- ulations still needs further research as also pointed out by Hoser. Additionally Hoser does not refer to a particular specimen from his type locality and hence the restriction is not valid. The genus Phoxophrys Hubrecht, 1881 is divided into three subgenera. In the introduction to the genus Hoser claims that “as there has never been a definition or diag- nosis of Pelturagonia” Mocquard, 1 890 he will “provide one herein for the first time”. Hoser’s diagnosis only com- prises two characters while that of Mocquard (1890) is written in French, and longer with several characters. To diagnose the genus Hoser uses approximately 20 lines, all of which are copied from Inger (1960: 221) and include a comparison to Japalura, the genus several Phoxophrys species belonged to until Inger’s revision. Hoser’s nominate subgenus is defined by a minimally rephrased diagnosis of Phoxophrys tuberculata Hubrecht, 1881 again taken from Inger (1960: 225, seven lines). The diagnosis of the subgenus to accommodate P. cephalum (Mocquard, 1 890) only comprises two lines with two char- acters (“presence of nuchal crest . . . and an absence of a supraciliary spine”). The last subgenus only contains P. spiniceps Smith, 1925. This is defined within seven lines, all copied but slightly rearranged from Inger (1960: 224-225). The next genus Hoser is concerned with comprises the lizards of the genus Aphaniotis Peters, 1864. The genus is divided into two subgenera on the basis of whether a “protrusion on the snouf ’ is present or absent. The genus and nominate subgenus diagnoses are identical and each constitute a copy from Boulenger (1885: 274, four lines). The other subgenus is defined by approximately four ad- ditional lines that have been copied from the internet (www.ecologyasia.com) or a source that we have not iden- tified. The genus Ptyctolaemus Peters, 1 864 currently consists of two species, which Hoser considers to be two subgen- era. The nominate subgenus containing P. gularis Peter, 1864 is initially defined within 15 lines copied from Schulte et al. (2004: 230) followed by a comparison to P. collicristatus Schulte & Vindum, 2004 (Schulte et al. 2004) taken from the same source (five lines). The defi- nition of the subgenus for P. collicristatus is precisely the other way round, i.e Hoser first uses the same five lines from the comparison betwee P. gularis and P. collicrista- tus to define the species and then the definition of the genus (all copied from Schulte et al. 2004: 230). The genus Salea Gray, 1 845 is currently considered to contain two species (see below) and one highly question- able species {S. gularis Blyth, 1854). In his introduction to the genus Hoser states that “neither the genus ... or the subgenus being properly defined to date . . . this is done herein for the first time”. He does however not present a single character to do so that has not been the result of a copying process from Boulenger (1 885). Hoser breaks up the genus into two subgenera based on the respective de- scriptions of S. horsfieldii Gray, 1835 and S. anamallayana (Beddome, 1878) taken from Boulenger (1885: 251-252, 312-314) with 36 lines (annotated as “modified from Boulenger” but actually constituting a verbatim copy) and 22 lines, respectively. For the latter species he resurrects its original name proposed by Beddome (1878). The last genus Hoser deals with in this part of the pa- per is Draco Linnaeus, 1758 which has been a matter of intensive morphological studies in the 1980s by Inger (1983) and Musters (1983). In recent years phylogenetic studies by McGuire & Alcala (2000), McGuire & Kiew Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK Critical review of Hoser’s publications on agamid lizards 125 (2001) and McGuire et al. (2007) completed the picture. Hoser mainly uses these phylogenetic results and the tree of Pyron et al. (2013) to divide the genus into nine sub- genera and copies their respective diagnoses from Boulenger (1885) or the morphology based publications mentioned before. Not a single new character is introduced by Hoser. Hoser’s general description of the genus is giv- en in 4.5 lines all taken from Boulenger (1885: 253). On- ly the phrase “much-produced” is replaced by “much-ex- panded”. The first new subgenus Hoser proposes serves to accom- modate members of the Draco lineatus group (minus D. lineatus which is placed in its own monotypic subgenus, see below). Initially Hoser copies seven lines from McGuire et al. (2007: 181) to define the group including a statement related to a statistical analysis. However, Hoser does neither use nor refer to a statistical method in his sec- tion on methods. Subsequently he produces four lines form the same source to define his subgenus further (McGuire et al, 2007: 181) followed by a short description of D. bi- maculatus Gunther, 1864 taken from Muster (1983: 40) to distinguish this species from his subgenus. The last part of Hoser’s diagnosis serves to separate D. lineatus Daudin, 1 802 from his proposed subgenus of the remaining linea- tus group species. This is done by copying the diagnosis comprising ten characters provided by McGuire et al. (2007: 199). At the end of this paragraph Hoser annotates “adapted from McGuire et al. (2007)” although he actu- ally produces a complete verbatim copy from that source. This goes so far that Hoser even has the typographical er- ror “posnuchal” [sic!] in the same place. The new monotypic subgenus to accommodate Draco bimaculatus initially repeats the four lines taken from Musters (s. above) followed by a copy (ten lines) from McGuire et al. (l.c.) as given under the previously defined subgenus. Next Hoser uses again the “adapted” diagno- sis for Z). lineatus provided by McGuire et al. (2007: 199, 16 lines including typographical error, see above) and fi- nally describes the species by copying Boulenger (1885:263, 19 lines) which again is annotated as having been “adapted” albeit constituting a word-for-word copy. Draco modiglianii Vinciguerra, 1 892 is placed by Hoser into its own new subgenus on the basis of a short diag- nosis (3.5 lines) that has been copied from Musters (1983: 45). Species related to Draco blanfordii Blanford, 1878 are combined in yet another new subgenus which he defines by copying three sets of characters originally from Boulenger (1885: 255, synopsis to the species, approxi- mately nine lines). No other characters are presented. Species related to Draco maculatus (Gray, 1845) are contained in a new subgenus that is entirely defined by 18 lines coming from Boulenger (1885: 262). The nom- inate subgenus is diagnosed in approximately three lines copied from Inger (1983: 17). Then Hoser defines Draco lineatus Daudin, 1802 in pretty much the same way he did to diagnose the linea- group (s. above). Initially he uses McGuire et al. (l.c.) to define the lineatus-group (approximately nine lines copied); this is followed by separating D. bimaculatus from that group and the proposed subgenus by copying Musters (1983: 40, 4.5 lines). Finally Hoser reproduces the full set of characters as given by McGuire et al. (2007: 199) for the species annotated as adapted but actually copied. For this subgenus Hoser resurrects an old avail- able name from Fitzinger (1843). Species related to Draco fimbriatus Kuhl, 1820 are placed into a subgenus for which another name proposed by Fitzinger (l.c.) is resurrected. The subgenus is defined in approximately three lines and subsequently separated from D. maculatus (again three lines), all copied from Boulenger (1885: 254-255). The last subgenus Hoser erects serves to accommodate the Indian species Draco dussumieri Dumeril & Bibron, 1837. To name the subgenus Hoser resurrects another of Fitzinger’s names (l.c.). The diagnosis consists of four lines taken from Boulenger ’s synopsis (1885: 255) fol- lowed by approximately 17 lines of description copied from the same source (Boulenger 1885: 268). After having defined his genera and subgenera Hoser endeavours to divide the subfamily into tribes and sub- tribes. Hoser proposes ten tribes and six subtribes, which will be numbered numerically in the following in order to prevent accidental validation; genus names are given here in their currently accepted form. Tribe 1 only contains lizards of the genus Draco. Tribe 2 contains the gmeraJapahira [in part] and Pseudocalotes (subtribe 2.1), Sitana and Otocryptis (subtribe 2.2), Acan- thosaura and Oriocalotes (subtribe 2.3) and Salea (sub- tribe 2.4). Tribe 3 only contains Calotes. Tribe 4 is rep- resented by Gonocephalus robinsonii and Japalura polygonata. Tribe 5 consists of Ceratophora, Cophotis, Pseudocophotis and Lyriocephalus (subtribe 5.1), Gono- cephalus mjobergi (subtribe 5.2), Gonocephalus, Bron- chocela, Complicitus, Hypsicalotes, Coryphophylax and Aphaniotis (subtribe 5.3). Tribe 6 comprises Japalura [in part, including Oriotiaris] and Ptyctolaemus. The remain- ing tribes contain a single genus each: Tribe 7 Lophocalotes, Tribe 8 Phoxophrys, Tribe 9 Mantheyus and Tribe 1 0 Dendragama. The nodes produced in Pyron et al. (2013) are given in the following as A-H with corresponding tribe numbers (as given above) from Hoser in brackets: A(l) - Draco, B(2) - Japalura Eastern clade, Pseudocalotes, Sitana, Otocryptis, Acanthosaura, Salea, C(3) - Calotes, D(4) - Japalura polygonata & Gonocephalus robinsonii, E(5) - Ceratophora, Cophotis, Lyriocephalus, Gonocephalus, Bronchocela, Coryphophylax, Aphaniotis, F(6) - Ptycto- laemus and Japalura variegata clade, G(8) - Phoxophrys and H(9) - Mantheyus. Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK 126 Wolfgang Denzer et al. As can be seen Hoser’s taxonomic scheme essentially reproduces the clades resulting from the phylogenetic analysis by Pyron et al. (2013). His subtribes can be de- rived in a similar way. Lophocalotes (Hoser’s Tribe 7) and Dendragama (Hoser’s Tribe 10) are not included in Py- ron’s analysis nor are Harpesaurus, Thaumatorhynchus, and Psammophilus. The first two are recognized by Hoser in their own monotypic tribes, the other three are not dealt with at all. Other genera also not included in Pyron et al. (l.c.) such as Oriocalotes, Hypsicalotes, and CompUcitus are assigned to a tribe but without giving a reason for do- ing so. However, with a bit of nomenclatural research one could find possible reasons for his groupings: Oriocalotes paulus was considered by Boulenger as Acanthosaura mi- nor, hence Hoser’s pairing of these two genera. In his in- troductory sentence to the genus CompUcitus he states that it was “formerly placed in Bronchoceld\ In the same pub- lication (Malkmus 1994, a paper written in German) Hyp- sicalotes is also considered to be a member of the genus Bronchocela. This is presumably Hoser’s reasoning be- hind grouping these two genera in the same tribe along with Bronchocela and several other species from the same node in Pyron et al. (l.c.). Had Hoser decided to follow the majority of earlier publications, all of which are cit- ed by him, he would likely have included these two gen- era in the tribe containing Calotes, their original genus name. Hoser’s division of the subfamily Draconinae can on- ly be understood and followed if Pyron’s paper is at hand for comparison. His classification scheme is poor-quali- ty if not worthless as most tribes and subtribes are not di- agnosed by shared characters but only through their con- tent. In Hoser’s words: ’’...tribe is best defined by diag- nosis of the component genera” or a similar wording. Such a definition may comprise the character sets of 13 genera as given in the first section of his paper where the genera are defined. All diagnoses are copied and no additional da- ta or characters are given. Some genera were not defined in their own right in Hoser’s first section of the paper. These genera are therefore diagnosed by him as a char- acter set defining a tribe or subtribe. In the following we will briefly analyze these additional diagnoses: Hypsicalotes Denzer & Manthey, 2000 is diagnosed by repeating entirely the diagnosis including comparisons to other genera as given in Denzer & Manthey (2000, ap- proximately 60 lines). Coryphophylax Fitzinger, 1869 is defined in the spaee of six lines copied from Boulenger (1885: 282). Cophotis Peters, 1861 is diagnosed by copy- ing Boulenger (1885: 251-252, three lines; 275, three lines). Pseudocophotis Manthey in Manthey & Gross- mann, 1997 is a copy of Boulenger ’s description of Cophotis sumatrana (= Pseudocophotis sumatrana). CompUcitus Manthey in Manthey & Grossmann, 1997 is defined in approximately two lines taken from the Rep- tile Database website (primary source not identified). Lophocalotes Gunther, 1 872 is diagnosed in approximate- ly eight lines of which four lines each are from Boulenger (1885: 251) and de RooiJ (1915: 116) or partially from Hallermann (2004). The genus Phoxophrys Hubrecht, 1881 is defined for a second time but this time using Boulenger (1885: 251, 280, six lines copied) instead of using Inger (1960). The definition of Dendragama Doha, 1 888 has been copied (approximately three lines) from de RooiJ (1915: 117-118). Summary Section A Apart from some minor alterations the phylogentic tree published by Pyron et al. (2013) serves as Hoser’s main basis for his classification scheme of the Draconinae. In our analysis of Hoser’s proposed taxonomy for the subfamily Draconinae most characters were identified and can be attributed to other sources. Hoser gives his diag- noses in the space of 2430 lines where 1884 lines consti- tute the actual characters of which 1560 lines have been identified as identical copies. If only diagnostic charac- ters are taken into account approximately 83% are a copy and if the full diagnoses are considered this percentage still comes to approximately 64%. If the full paper is taken in- to account (7140 lines, estimated) the copied text still amounts to approximately 22%. At the end of his paper Hoser cites several hundred references in a space of close to 16 pages. However, according to our analysis he only used approximately 30 of those to produce his proposed taxonomy, the bibliography of which could probably have been printed in the space of two pages. If this is taken in- to account the percentage of copied text in relation to the full paper rises by yet another 5%. Hoser’s main source for descriptive characters was Boulenger (1885). Additionally he copied sections from Inger (1960) for Phoxophrys, Denzer & Manthey (2000) for Hypsicalotes, Hallermann (2004) for Bronchocela, McGuire et al. (2007) for Draco, Denzer & Manthey (2009) for Gonocephalus, Mahony (2009) for Japalura, Zug et al. (2006), Krishnan (2008) and Hallermann (2000) for Calotes, Harikrishnan & Vasudevan (2013) for Pseudocalotes, Pethiyagoda & Manamendra-Arachchi (1998) for Ceratophora and Schulte et al. (2004) fox Ptyc- tolaemus. Several genera such as Harpesaurus, Thauma- torhynchus, and Psammophilus are not treated at all. B) Hoser (2013) on Amphibolurinae We note that the manuscript on Amphibolurinae was re- ceived hy AJHoxv 20 July 2013, accepted for publication on 4 October 2013, and published on 20 October 2013. However, a tax invoice printed at the end of the publica- tion (p. 36) states that the journal was printed on 3 Octo- ber 2013, implying printed copies may have existed be- fore the paper was accepted. Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK Critical review of Hoser’s publications on agamid lizards 127 The paper comprises the following sections or headings: Title, Abstract (including Keywords), Introduction, fol- lowed by the description of two genera and seven tribes, References Cited, and a statement about Conflict of Inter- est. An explicit section for Materials is missing (but see below) and one has to assume that the description of the newtaxa constitutes a combined Results/Discussion/Con- clusion section. The introductory part of a publication typically includes a brief overview and often the author’s motivation for writ- ing the paper, as well as his ideas about the subject. In stan- dard practice, reviewers of manuscripts in mainstream journals would not spend much effort in correcting this part unless false statements are presented. However, in the case of this particular introduction, it is instructive for a better understanding of the broader picture of Hoser’s works to mention several paragraphs. Hoser begins by providing a reason for why the Aus- tralasian Amphibolurinae Wagler, 1830 are so well stud- ied. According to Hoser this “has arisen due to a combi- nation of circumstances relatively unusual to Australia” (emphasis added). The two factors alluded to are a “sta- ble political and economic situation” including a transport infrastructure that facilitates access to even the most re- mote parts and “well-funded government paid herpetolo- gists and relatively wealthy . . . private herpetologists . . . able to travel to the most remote parts of the continent. . . A significant portion of the Introduction deals with the publications by Wells and Wellington (1983, 1985), which are considered highly controversial papers in their own right and still do not find full acceptance within the her- petological community. One part of a paragraph reads as follows (Hoser 2013: 34): “I have found myself resurrect- ing names proposed by earlier authors. This includes a number of effectively unused Wells and Wellington names such as IntellagamaWQWs & Wellington, 1985, Gowidon Wells & Wellington, 1 983 ...” However, at the time of this paper’s publication (October 2013) the genus name Intel- lagama had already been validated by Amey et al. (2012) and the genus name Gowidon in the combination G. lon- girostris (Boulenger, 1883) was made available by Melville et al. (2011). Both publications were not cited by Hoser (2013). One of the paragraphs in Hoser’s Introduction would never pass standard review of any formal publication in science and would be removed by editors as it is against ethical standards of publication. A group of herpetologists (one named in particular) that is highly critical of Hoser’s papers is called “a mob of criminals and ratbags” (Hoser 2013: 34). At the end of the Introduction, Hoser uncritically lists 14 publications, five of which are his own, “and sources cited therein” that apparently constitute the source mate- rial for his research. However, major publications on Aus- tralian agamids relevant to taxonomy and nomenclature. such as Melville et al. (2011), Hugall & Lee (2004), Hugall et al. (2008), Schulte et al. (2003), or Macey et al. (2000a,b) are not mentioned at all and the reader would need to refer to the few cited papers and their bibliogra- phies to determine how Hoser derived some of his ideas. While this paragraph could be regarded as a Materials sec- tion, it should be noted that not a single museum speci- men is referenced, nor is there any mention that museum material was examined. Earlier in the Introduction Hoser (2013: 33) notes: “In terms of the materials and methods, this was based on my own field and lab work involving most species as well as a review of the relevant literature spanning the last 200 years.” With respect to “lab work,” the reader does not get any further explanation of what this entailed, making the process non-transparent and non- reproducible. The list of references given in the References Cited section of the paper comprises only 16 citations, five of which are Hoser’s own publications. Of those, at least Hoser (1998) on Acanthophis and Hoser (2012) on Afron- aja are entirely irrelevant to agamid lizard taxonomy. Only Joger (1991) and Pyron et al. (2013) are referenced as publications that include original molecular phyloge- netic research that is indispensible to Hoser’s arguments. However, Joger ’s paper (l.c.) only includes Amphibolu- rus vitticeps (Ahl, 1926) (= Pogona vitticeps) and Physignathus temporalis (Gunther, 1867) (= Lophog- nathus temporalis) in the analysis, making it of only pe- ripheral interest for a detailed phylogenetic analysis of Australian taxa (Joger 1991, Material Examined). Joger (1991: 619) even notes: “Because of the lack of antisera for East Asian and Australian agamids, the position of their three lineages - AmphibolurusI Physignathus, CaloteslAcanthosaura, and Gonocephalus - relative to each other could not be determined.” It is important to note that Joger did not study the genera Physignathus Cuvier, 1829 mdAmphibolurus Waglor, 1830, but two specimens of populations that were considered members of these gen- era at that time, but are assigned to different genera to- day. Additionally, no nomenclatural decisions were pro- posed by Joger (l.c.). It therefore stands to reason that Pyron et al. (2013) serves as the basis for Hoser’s taxonomic and nomenclat- ural proposals. This becomes particularly obvious in the grouping of Moloch Gray, 1841 and Chelosania Gray, 1 845 within a single tribe. Without recent molecular phy- logenies it is unlikely that any morphologically-oriented herpetologist would group a thorny devil (whose vernac- ular name illustrates a key aspect of the species’ scale mor- phology) with a lizard that has a completely homogeneous dorsal scalation. The phylogeny of Pyron et al. (2013) is also reflected in acknowledging the difference between Physignathus and Intellagama, the splitting of Hypsilu- rus into several genera, the erection of a tribe for the genus Ctenophorus, and combining Amphibolurus, Chlamy- dosaurus, Diporiphora, Gowidon (Lophognathus), Pog- Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK 128 Wolfgang Denzer et al. ona, Rankinia, and Tympanocryptis within one tribe, al- beit using a different nomenclature. The Introduction is followed by a section that serves to erect two new genera and seven tribes. New tribes are pre- sented in random order without presenting necessary in- formation on possible phylogenetic relationships. Our analysis will mainly follow Hoser’s order but for reasons of clarity the tribe containing the genera related to Am- phibolurus and the tribe containing all Ctenophorus will be dealt with last. The first genus described in the paper is meant to ac- commodate only Diporiphora superba Storr, 1974. The actual diagnostic characters account for approximately nine lines of text, of which eight can be accounted for in Cogger (1983: 238, key leading to D. superba, 1983: 243, description of D. superba). Differences include changing “gular fold absent” to “no gular fold” and replacing the % sign by “percent,” as well as replacing the numeric “4” by the word “four.” The unaccounted text consists main- ly of introduced verbs and a slightly modified description of the colouration. The second genus described deals with Hypsilurus spinipes (Dumeril & Bibron, 1851). The diagnostic char- acter section accounts for approximately 12 lines and is annotated as “adapted from Cogger, 2000.” About 11 lines are the result of directly repeating Cogger’s description of H. spinipes (Cogger 1983: 245-46, as Gonocephalus spinipes) and the only differences are the introduction of a few verbs and conjunctions. The new genus containing Hypsilurus spinipes is sub- sequently placed into a newly erected tribe that addition- ally contains Tiaris Dumeril & Bibron, 1837 (see below). The diagnosis for the tribe contains two sets of charac- ters. The first part (ca. seven lines) is a copy of Cogger’s key leading to Hypsilurus (Cogger 1983: 217, as Gono- cephalus) apart from a few introduced verbs and conjunc- tions (one line). The second part (eight lines) states the characters shared by Hypsilurus species and is a copy (six lines) of the genus diagnosis given by Manthey & Denz- er (2006). The latter paper is not cited, and the diagnosis was most probably retrieved from the Reptile Database (Uetz & Hosek 2015), where it is publicly available (cit- ed, and with approval of the authors). The genus Tiaris is not characterized at all in Hoser’s paper, nor is the read- er informed which species it contains. In Hoser’s paper the name stands on its own and is therefore a nomen nudum according to the Code. It should also be noted that Tiaris Dumeril & Bibron, 1837 is not available for any agamid genus as it is preoccupied by Tiaris Swainson, 1 827 (Aves, Passeriformes) [see Manthey & Denzer (2006 )]• According to Hoser an agamid genus Tiaris is of Aus- tralian origin and “the only genus it is likely to be con- fused with” is the one newly erected for Hypsilurus spinipes. The only other Hypsilurus species in Australia that Hoser could refer to is H. boydii (MacLeay, 1884), albeit that it is actually quite difficult to confuse these two species. Hypsilurus boydii is known from older literature as Tiaris boydii (e.g., MacLeay 1884). Hence, we assume that Hoser meant to include this species in the same tribe as H. spinipes. However, H. boydii is morphologically (Manthey & Denzer 2006) and genetically (Pyron et al. 2013) closely related to H. dilophus (Dumeril & Bibron, 1837), which Hoser places into a different tribe. The next tribe is erected to accommodate the genus Hyp- silurus, assigning Lophura (Hypsilurus) godeffroyi Peters, 1867 as the terminal taxon. Hoser’s introduction to the tribe starts with the sentence “Currently most widely known as Hypsilurus dilophus (Dumeril & Bibron, 1837).” It is not clear whether Hoser here expresses his view that H. godeffroyi (a well-defined and valid species) is identical to (conspecific with) H dilophus. His state- ment is even more confusing considering that H. dilophus is actually the type species of the genus Tiaris Dumeril & Bibron, 1 837, a genus he assigned a paragraph earlier to a different tribe (for synonymy of Hypsilurus see Man- they & Denzer 2006). Additionally there exist no objec- tive reasons to combine H dilophus and H. godeffroyi in- to one group. Morphologically, they are very different species that were even placed into different species groups by Manthey & Denzer (2006). In his genus description to accommodate H. spinipes, Hoser states that Tiaris (nomen nudum, see above) and all Hypsilurus species have a “lon- gitudinal row of grossly enlarged scales on the throat.” This is actually a character used by Cogger (1983) to dif- ferentiate between H spinipes and H boydii, which should read, “median longitudinal line of . . . similar to those in the nuchal crest.” The statement holds true if only Aus- tralian species are considered, as is the case with Cogger (I.C.), but when including taxa outside of Australia, as Hoser’s analysis does, it is false, as most species of Hyp- silurus outside Australia actually lack a median line of en- larged scales on the gular pouch. These are only well de- veloped in H. boydii and H. dilophus, and to a lesser ex- tent in H. hikidanus Manthey & Denzer, 2006. As already pointed out by Manthey & Denzer (2006), H dilophus, H boydii, and H. spinipes may be considered as a species group, and if considered as a separate genus only the name Lophosaurus Fitzinger, 1843 would be available but not Tiaris. The diagnostic character section for Hypsilurus comprises 15 lines of which 13 lines are copied from Cog- ger (l.c.) and Manthey & Denzer (2006). Subsequently, Hoser deals with the water dragons from Australia and Southeast Asia. The relationship, biogeog- raphy, and nomenclature of Physignathus cocincinus Cu- vier, 1 829 and Intellagama lesueurii (Gray, 1831) has been a matter of intense discussion, and since the advent of mo- lecular phylogeny there have been several publications to address the issues (e.g., Schulte et al. 2003, Macey et al. 2000a,b). Still Hoser makes no mention of this and erects Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK Critical review of Hoser’s publications on agamid lizards 129 two new tribes. His claim to have resurrected the genus name Intellagama has already been dealt with (see above). The diagnostic characters to define the tribe for 7. lesueurii account for ca. five lines of text, three of which are copied from Cogger’s key (Cogger 1983: 217). The remaining part of the description deals with colouration, but not as one would expect, with a description of the colouration of 7 lesueurii. Hoser instead merely states that the coloura- tion is not that of P. cocincinus. The tribe containing the latter species is again initially diagnosed via Cogger’s key (f c.), with three lines out of four being copied. The remaining part of the diagnosis deals with the colouration of Physignathus cocincinus. Apart from the dorsal ground colour (one line) the pat- terns and colourations (three lines) were copied from Wikipedia, with no primary source identifiable. The Wikipedia page already provided this description of the colouration in 2012 (accessed August 2014, file history checked for December 2012), i.e. before Hoser’s paper was published. Next Hoser introduces a tribe to accommodate the gen- era Moloch and Chelosania. The two diagnostic sections consist of two lines and four lines, respectively, of which approximately four lines are copied text (Cogger 1983: 217). Another tribe is erected to accommodate the genus Ctenophorus. The diagnostic characters are presented in roughly sixteen lines, ten of which are copied from Cog- ger (1983) and three from Cogger (1993). One set of char- acters regarding the supralabial scales could not be ac- counted for and is potentially the only part of an original description in the entire paper. From the lack of method- ology, it is not possible to determine how these observa- tions were made or which specimens were used, render- ing the data non-reproducible. Furthermore, the copied part contains several mistakes that need to be addressed. The diagnosis for the new tribe containing the genus Ctenophorus is at best confusing, and perhaps of no tax- onomic value entirely. Part of Hoser’s definition reads as follows (emphasis added): “nuchal crest and/or series of enlarged keeled vertebral scales present or absent and if absent present along at least the anterior two thirds of the body; enlarged strongly keeled or spinose scales are present elsewhere on the dorsum.” The diagnostic char- acters are identical those used in Cogger’s key to the gen- era (1983: 217), apart from the conflicting phrase “absent present” and the placement of the semicolon. The use of both “absent” and “presenf ’ in close combination makes it unclear how this character is to be scored. In common usage, placing a semicolon will not change a diagnosis sig- nificantly. However, in this case only the part directly pre- ceding the semicolon relates to the character of “enlarged keeled vertebral scales present or absenf’. The character after the semicolon “enlarged strongly keeled scales . . . present ... on the dorsum” stands on its own. This way all Ctenophorus without this character are excluded from the genus! Cogger (1983: 217) included this particular set of characters as a full statement in the diagnosis for the genus Amphibolurus. There it reads, ’’nuchal crest and/or vertebral keel may be present, but if the latter is present on at least the anterior two-thirds of the body then en- larged, strongly-keeled or spinose scales are present else- where on the dorsum”, giving it a completely different meaning. It should be noted that a key matching this part of Hoser’s diagnosis, including the (wrong) placement of the semicolon, can be found in another earlier publication (Cogger 1993: 163, or in the online version on page 10, character 9a). A further character to define the tribe pre- sented by Hoser is described on the basis of the online pub- lication (page 11) but introducing yet another mistake. Hoser’s character reads: “a row of enlarged scales from below the eye to above the eye” instead of “to above the ear”! The genus- and species-richest tribe introduced in the paper contains the genera Amphibolurus, Chlamy- dosaurus, Caimanops, Cryptagama, Diporiphora, Gowidon (Lophognathus), Pogona, Rankinia, and Tymap- nocryptis, as currently accepted by most Australian her- petologists (here listed according to Cogger 2014; it should be noted that Gowidon is not yet generally accepted). However, according to Hoser’s compilation the genus Lophognathus no longer exists. Hoser instead uses Gowidon, a name available for 7. longirostris, but ignores 7. Wells & Wellington, 1985, 7. gilberti Gray, 1842, and 7. temporalis (Gunther, 1867). For this reason the reader has to assume that Hoser considers these species as congeneric or even conspecific. However, in this case the name Gowidon would not be available since the genus name Lophognathus has nomenclatural priority over it, with 7. gilberti being the type species of the genus. If Hoser had been consistent in following the data of Py- ron et al. (2013), then Tympanocryptis Peters, 1 863 should also be a member of this tribe. However, Hoser does not include it here or in any other tribe, nor does he use a dif- ferent taxonomy to pinpoint where the species of this genus might be grouped, perhaps as part of one of the oth- er genera used in the revised classification scheme. It ap- pears that, just as some members of Lophognathus, the genus Tympanocryptis was simply disregarded or forgot- ten. Tympanocryptis is an available name that should be used, the type species being T lineata Peters, 1863. Hoser recognizes Caimanops Storr, 1974 and two gen- era proposed by Wells & Wellington (1983, 1985) con- taining Diporiphora species. One of these genera was erected to accommodate D. albilabris albilabris Storr, 1974 and 7). albilabris sorbia Storr, 1974. The second was erected for 7). /zwga Houston, 1977 and 7). winneckei Lu- cas & Frost, 1 896. We assume that Hoser resurrects these genera from their synonymy with Diporiphora, albeit without mentioning it specifically or giving a reason for Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK 130 Wolfgang Denzer et al. doing so. These genera were formally synonymized with Diporiphora by Doughty et al. (2012), owing to their close phylogenetic relationship, and were included by Pyron et al. (l.c.) under Diporiphora. It appears that Hoser over- looked the publication by Doughty et al. (2012). Caimanops has so far been considered a monotypic genus, with C. amphiboluroides as the sole species. If Hoser had properly followed Pyron et al. (l.c.), he should also have assigned D. australis to Caimanops as these two species form a clade. The diagnostic characters employed by Hoser to group the above genera into a new tribe are as follows: Initial- ly he singles out Chlamydosaurus by repeating Cogger (1983 two out of two lines copied) but introducing a mis- take. Hoser ’s character reads, “a large loose frill or skin around the neck” but it should read, “. . . frill of skin . . . ”. Hoser then proceeds to define the general characters of this diverse group (Cogger 1983; four out of four lines copied). In his last part of defining the tribe he excludes the genus Ctenophorus in an identical way as he defined the tribe containing the genus, including all the mistakes discussed above. Overall Hoser’s diagnostic character sec- tion of this tribe comes to 22 lines of which 16 lines are a result of copying. Summary Section B Hoser’s classification scheme for amphibolurine lizards mostly reflects the nodes in the phylogenetic tree pub- lished by Pyron et al. (2013). Additionally several genera proposed in the highly controversial papers by Wells & Wellington (1983, 1985) are accepted as valid. Hoser gives his diagnoses in the space of 153 lines, of which 121 lines constitute the actual characters, with 100 lines clearly identifiable as copied. If only diagnostic char- acters are taken into account this amounts to 82% copied material, with the full diagnoses included it is still 65%. If the full paper is taken into account (438 lines) the copied text amounts to 23%. All but a single character can be identified and attrib- uted to secondary sources (Cogger 1983, 2000; Manthey & Denzer 2006; Anonymous on Wikipedia; Uetz & Hosek 2015). Most characters used to describe genera are taken di- rectly from Cogger (1983 or subsequent editions). Sever- al important publications on Australian agamid lizards such as Melville et al. (2011), Hugall & Lee (2004), Hugall et al. (2008), Schulte et al. (2003), or Macey et al. (2000a,b) have been omitted. The genera Lophognathus and Tympanocryptis are not treated at all. C) Hoser (2012a) on Laudakia Gray, 1845 The manuscript was received on 13 March 2012, accept- ed on 8 April 2012 and the paper was published on 30 June 2012. Hoser’s paper is presented in the following way: Ab- stract (including Keywords), Introduction, description of taxa, and References Cited. The second to fifth paragraphs of the Introduction, describing the general appearance and behaviour of the group, contain 13 lines of copied mate- rial from an online source (www.sauria.co.uk). The rest of the Introduction deals with the nomenclature and phy- logeny of Laudakia Gray, 1 845 and Phrynocephalus Kaup, 1825. His nomenclatural arguments refer mainly to Hen- le (1995) and are misinterpreted (see comments for the stellio group). Hoser’s “five- way division” of Laudakia Gray, 1845 mainly reflects the phylogenetic schemes published by Macey et al. (1998, 2000b, 2006), who identified nodes supporting a L. tuberciilata group, a L. caucasia group with L. lehmanni as the sister taxon (proposed as a new subgenus by Hoser), as well as nodes supporting the monophyly of L. stellio (Linnaeus, 1758) and L. sacra (Smith, 1935). Hoser’s fifth group comprises Phryno- cephalus Kaup, 1825, a genus of lizards that has never been in the synonymy of Laudakia. Macey et al. (2000b) found Phrynocephalus to be a sister taxon to both the clade containing the L. caucasia group and L. stellio. Howev- er, in a later publication by Melville et al. (2009), the monophyly of the genus Laudakia was confirmed and Phrynocephalus emerged as the sister taxon to the whole clade. Apparently, Hoser and his supposed reviewers over- looked this important publication, which is not cited in his bibliography. The first genus Hoser deals with is that of Phryno- cephalus Kaup, 1 825, which he considers “similar in most respects to Laudakia sensu lato” (Hoser 2012: 1 8), a state- ment most herpetologists would disagree with. Hoser does not present a meaningful definition of the genus apart from “lacking of an obvious tympanum” to distinguish Phryno- cephalus from Laudakia and a “dorsoventrally de- pressed” body to distinguish it “from all other other Agamids in the region where these groups of lizards oc- cur.” This entire diagnosis holds no definitive value as there are other agamid genera in the area under consider- ation that have a dorsoventrally depressed body shape (e.g., Brachysaura Blyth, 1856, Bufoniceps Arnold, 1992, and Trapelus Cuvier, 1829). Another interesting fact is that Hoser only recognizes 26 species within the genus Phrynocephalus while the actual number had already sur- passed 40 species by the time his paper was published. This is certainly something any expert reviewer would have been able to point out, even by a simple search of the Reptile Database (Uetz & Hosek 2015). Hoser moves on to define what he considers to be the actual genus Laudakia. This is represented by the tuber- Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK Critical review of Hoser’s publications on agamid lizards 131 culata group and hence L. tuberculata (Gray, 1 827) be- comes the type species. Initially he uses tympanum pres- ence and body shape to distinguish it from Phryno- cephalus and other genera of agamid lizards in the region. This is followed by a short description (eight lines) most- ly copied from a key to the species of agamid lizards of Pakistan (six lines; Khan 2002: 100 & 101). The same key is available on the Internet as part of Khan’s undated eBook Herpetology of Pakistan. Neither of these two pub- lications are cited by Hoser! Based on the idiosyncratic character “fifth toe extends beyond second” we have no doubts that Khan’s publication is the source; other authors make a comparison to the first toe. Hoser’s further char- acters include the dentition and the number of whorls in each tail segment. Both characters are from Baig (1992) but the writing has been sufficiently altered that they do not constitute copied material. The entire diagnosis of the genus comprises 32 lines (18 lines of diagnostic charac- ters) of which six lines are copied, presumably from Khan (2002). The next genus proposed by Hoser serves to accommo- date the Laudakia caucasia (now Paralaudakia caucasia, see below) group. Up to this point we have mostly ab- stained from judging Hoser’s diagnoses but the poor qual- ity of this one requires analysis. It begins with an essen- tially copied general diagnosis from earlier in the paper, but “excluding those genera formerly placed within Lau- dakia sensu lato” by which Hoser means the other gen- era he proposed (see above). Subsequently, he copies from Khan (2002): “tympanum is large,. . . fifth toe extends be- yond second; caudal scales in distinct annuli,” which is unfortunately a character set that still defines the entire genus. Up to this point, no character has been listed that could be used to define the new genus. Next, Hoser states that “the scales of dorsal rows are smooth,” “the premax- illa has two teeth in the [new] genus versus three in Lau- dakia [= L. tuberculata group],” and “lizards in this [new] genus have 14-15 molars, versus 14-15 [sic!] in Lau- dakia.'" The author then once more repeats the general paragraph to differentiate Phrynocephalus and other agamids. He then separates his new genus from L. sacra by providing a full description of this species that is iden- tical to the one found in Ananjeva et al. (1990). To sum- marize this for clarity: the only diagnostic characters pre- sented to define his new genus -other than those charac- ters which are common to all genera concerned- are “scales of dorsal rows are smooth,” “premaxilla has two teeth,” and presumably the number of molar teeth. The definition for the new genus is, unfortunately for Hoser, not cohesive because the vertebral scales of Par- alaudakia caucasia (Eichwald, 1831), the proposed type species of the new genus, are actually keeled and those of P. himalayana (Steindachner, 1867) and P. badakhshana (Anderson & Leviton, 1969) are smooth (Boulenger 1885; Khan 2002; Baig et al. 2012). We are not aware - and it is outside the scope of this paper to in- vestigate further - how many teeth are present in the pre- maxilla and how many molars the other genera possess, in order to verify or falsify these two characters, nor is Hoser apparently. To our knowledge there are no publi- cations dealing with the dentition of all genera in ques- tion. In total his diagnosis of this genus comprises about 56 lines, of which 27 are copied from Ananjeva et al. (1990). Three lines describing diagnostic characters are taken from Baig (1992) and Khan (2002), but not copied directly. Baig et al. (2012) established the genus Paralau- dakia Baig, Wagner, Ananjeva & Bohme, 2012 to accom- modate species related to caucasia and himalayana as well as lehmanni and stoliczkana. For the Laudakia stellio (now Stellagama stellio, see be- low) species group, which Hoser considers to be mono- typic, the author resurrects Plocederma Blyth, 1 854. This can only be explained by misinterpreting Henle (1995) who proposed to use this genus name for the stellio group, which he considered to comprise L. stellio, L. caucasia, L. erythrogastra, L. himalayana, L. lehmanni, L. nupta, and L. melanura. The type species for the genus Ploced- erma is L. melanura. Only if this species were included in Hoser’s stellio group (which it is not) would the name be available for the group. Because the stellio group as it is considered nowadays (i.e. monotypic) did not have any previous available name disposable, Baig et al. (2012) es- tablished the name Stellagama Baig, Wagner, Ananjeva & Bohme, 2012. The new genus to accommodate Stellagama stellio is initially only defined by repeating his general description (two characters: “distinct tympanum” and “dorsoventral- ly depressed” body) followed by two lines taken from Khan (2002) and a description of ''Laudakia stellio" (ap- proximately 14 lines, all copied) taken from a website (Go 9 men, www.bayramgocmen.com/album/picture.php? /1012/category/345, accessed September 2014). The full diagnosis comprises 26 lines (18 lines of diagnostic char- acters), of which 16 are copied. To define his newly proposed monotypic subgenus to accommodate Laudakia lehmanni (Nikolsky, 1896), Hoser presents diagnostic characters in the space of ap- proximately 38 lines, all of which come straight from Baig’s description (1992) of L. lehmanni. The order of characters is slightly different from the original and in a few places verbs or conjunctions have been added. The diagnosis is followed by distributional data and habitat de- scription, which constitutes (apart from one sentence) a copy of the text produced on the lUCN RedList webpage (six lines out of seven copied). The whole diagnosis com- prises 42 lines of which 38 lines constitute diagnostic char- acters all of which are a copy from Baig (1992). Hoser lists four papers by Baig and co-authors in the References Cited section, but none of these contains a de- scription of L. lehmanni (now Paralaudakia lehmanni, see Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK 132 Wolfgang Denzer et al. below). A detailed description of P. lehmanni that is very similar to the one used by Hoser - but with characters in a different order - was first given in Baig’s PhD thesis (Baig, 1992: 130 & 132, not cited by Hoser) which was completed under the supervision of WB. Since Baig’s the- sis has to be considered unpublished, Bohme and cowork- ers (Baig et al. 2012) posthumously published a paper based on the thesis to preserve Baig’s extensive taxonom- ic work and to make it available for scientists working on this subject. In the latter publication Baig’s description is repeated with minor changes and with pretty much the same wording seen in Hoser (2012a). Baig et al. (2012) was published in print on 1 8 July 2012 and Hoser (2012a) was published in print 30 June 2012. Both papers were accepted for publication by the respective journals in April 2012. We also note that Baig et al. (2012) was made avail- able in advance online on the publisher’s website on 6 Ju- ly 2012, appearing a week after Hoser’s publication. The most likely way by which Hoser would have been able to retrieve Baig’s text would have been by downloading the thesis from a governmental website in Pakistan (Pakistan Research Repository, http://eprints.hec. gov.pk/2407/1/ 2262.htm). Although it is difficult to proof, but based on the exact wording, we are convinced that Baig’s thesis was available to Hoser, who did not consider it necessary to reference it. However, even if not published a PhD the- sis constitutes intellectual property belonging to the can- didate and his thesis supervisor. In any case the precise repetition of wording from a thesis without appropriate clarification, attribution and referencing constitutes a vi- olation of authorship rights. Any use of a verbatim copy of excerpts from a thesis needs permission by either the author, his thesis supervisor or the university department where the candidate studied for the degree. However, a reader who does not know about Baig’s thesis might sup- pose that Hoser’s diagnosis has precedence, with Baig et al. (2012) copying Hoser’s ideas and wording when the opposite is the case. In this instance, Hoser clearly uses the intellectual property of another and passes it off as his own. Such behaviour would even be seen as plagiarism if Hoser obtained the description from a third source, which we have not identified. In a recent paper Hoser (2015) even claims priority and that “they [Baig et al. (2012)] did however remanufacture theirs [morphologi- cal evidence] as “new” data, which in itself is fraudulenf ’. Hoser (l.c.) clearly states that data were available from ear- lier studies but again does not disclose or cite the source. Not only did Hoser plagiarize Baig (1992), he even con- siders his actions as justifiable and additionally accuses the true original author of fraudulent behaviour! The last genus Hoser proposes is monotypic for Lau- dakia sacra (Smith, 1935). His diagnosis is given within 27 lines, of which 25 are a direct copy from the descrip- tion of L. sacra by Ananjeva et al. (1990; see also Uetz & Hosek 2015) and two lines are copied from Khan (2002). The only two other characters are those used pre- viously (“a distinct tympanum” and “the body is dorso- laterally depressed”), repeating nearly the entire general paragraph for the fifth time. The diagnosis of this genus is given in 33 lines, with 30 lines presenting the actual di- agnostic characters, of which 27 lines are copied. Summary Section C The taxonomic basis for Hoser’s proposals on Laudakia can be found in their entirety in Macey et al. (1998, 2000b, 2006). Most of Hoser’s proposed classification addition- ally reflects nodes in the phylogeny published by Pyron etal. (2013). In total, Hoser’s paper on Laudakia comprises an esti- mated 980 lines, of which 420 lines constitute his Refer- ences Cited section (560 lines pure text including Title and Abstract). We would like to mention that already his in- troductory part contains at least 13 lines that can be found on websites (not taken into account here as copied text) and that we further identified several diagnostic charac- ters Hoser used but without copying directly. His diag- noses come to 180 lines of which 148 lines constitute di- agnostic characters. With respect to the latter we found that 114 lines (77%) were copied from previously published research papers or reviews. Hoser’s main sources are Ananjeva et al. (1990), Baig (1992), Khan (2002) and Baig et al. (2012). Hoser (2015) even claims priority with re- spect to the data albeit that his taxonomic scheme and all his characters have been copied from Baig (1992) and sub- sequent publications. D) Hoser (2014c) on Uromastycinae The manuscript of this paper was received by the journal on 2 November 2013, accepted on 15 May 2014, and fi- nally published on 30 August 2014. It is presented in the following way: Abstract (including Keywords), Introduc- tion, Notes on the taxa named herein followed by the de- scription of taxa. Conflict of Interest, and References Cit- ed. In total Hoser newly describes or resurrects within this publication two tribes, five genera, and four subgenera. Five of these taxa are monotypic. The Introduction is relatively short and summarizes the taxonomic history of Uromastyx and gives Hoser’s view on taxonomy, without any identified copied parts. How- ever, two extraordinary statements should be discussed here. In terms of material used for his study, Hoser refers to the “inspection of live specimens at various facilities since 1993.” Many Saara or Uromastyx species inhabit political unstable areas and it is very unlikely that there are live specimens of many important species, such as S. asmussi (Strauch, 1863), available at any facility Hoser might have visited since 1993. Therefore it is very prob- lable that most of the data he presents are not from ex- Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK Critical review of Hoser’s publications on agamid lizards 133 amined material, but from published sources. These sources are cited in the Introduction as “Significant stud- ies relevant to the taxonomy of the Uromastycinae ...” and include for example Hall (1999) and Swofford (2002), two general publications about statistical phylogenetic meth- ods without any significance to the taxonomy of the group at all. In his Abstract and Introduction, Hoser gives the im- pression that he is the first to use the approach by Pyron et al. (2013) to distinguish between Uromastycinae Theobald, 1868 and Leiolepidinae Fitzinger, 1843 on the subfamily level, but mentioning in passing that “some au- thors have already taken this step.” This concept was used more than a decade ago by Macey et al. (2000a), in a pa- per not cited by Hoser. Instead, Hoser cites Macey et al. (2000b) on the trans-Tethys migration, which has hardly any relevance to uromasticine / leiolepidine taxonomy (on- ly one species of Uromastyx and two species of Leiolepis were included in the study). We also note that the terms Uromastycinae and Leiolepidinae were used synonymous- ly by different authors (e.g., Wilms & Bohme 2007: 436). The entire first definition of Uromastycinae used by Hoser is identically phrased to Wilms et al. (2009:67; four lines), followed by the definitions of his two new tribes copied from the same source. The further detailed defini- tion for this subfamily is not fully copied, but obviously taken from Boulenger’s synopsis (Boulenger 1885: 405; 14 lines). All diagnosing parts of the subsequent defini- tion of Uromastyx are entirely taken from various parts (text and key) of Wilms et al. (2009). The first taxon Hoser describes is the subgenus Uro- mastyx (within Uromastyx) and the given diagnosis is tak- en from Wilms et al. (2009: 67 & 82; 35 lines copied for the genus Uromastyx, seven for the subgenus Uromastyx, of which 3.5 lines are within quotation marks). For his sec- ond, monotypic subgenus diagnosis, erected to accommo- date U. occidentalis Mateo, Geniez, Lopez-Jurado & Bons, 1999, he uses eight lines directly copied from Wilms et al. (I.C.). Even though Hoser quotes Wilms et al. (l.c., 3.5 lines) in the following diagnosis of Aporoscelis, the parts not di- rectly quoted are also copied from that reference (two lines). For his first new genus description Hoser is using a dif- ferential diagnosis separating his new taxon by the diag- nosis of other taxa. Here, Hoser is summarizing the in- formation given in the diagnostic key by Wilms et al. (2009: 82), followed by a description taken from Boulenger (1885: 405) with 26 lines copied. Also the di- agnosis of the same taxon at a different rank (subgenus) is taken from Wilms et al. (2009, 11 lines copied). With- in his new genus, Hoser describes two additional new sub- genera. Even here all mentioned characters diagnosing these taxa are identical to Wilms et al. (2009: 82; 22 lines copied). Diagnosing his second new genus, Hoser is following the same scheme of presenting a differential diagnosis. And again, all given characters, especially meristic char- acters (e.g., scale or whorl counts) are exactly the same as given in Wilms et al. (l.c.) and no other additional char- acters are provided. This description is followed by the subgenera to be included in the previously described genus. Again, all mentioned characters are exclusively tak- en from Wilms et al. (l.c.). The new genus is described by copying 14 lines from Wilms et al. (2009: 82-83), and subsequently three new subgenera are proposed using 44 lines from the same source. While redescribing the genus Saara according to his new taxonomy, Hoser provides several characters to dis- tinguish his new tribe, including Saara, from the tribe that includes Uromastyx. Here he is exclusively using the char- acters provided by Wilms et al. (2009: 81-82; 15 lines) for the three species forming the genus Saara. However, Hoser is splitting this genus into three distinct monotyp- ic genera, including in addition to Saara the genus Cen- trotrachelus Strauch, 1 863, which he resurrects to accom- modate S. asmussi, and a new genus that only contains S. loricatus Blanford, 1874. In order to describe these two genera Hoser again uses Wilms et al. (2009: 81-82; 32 lines copied). Finally, Hoser erects two new tribes to accommodate his proposed genera. The first tribe is described using three lines from Wilms et al. (2009:67) and 13 lines from Boulenger (1885:405). The second tribe is solely defined by characters given by Wilms et al. (2009: 67, 82 & 83; 15 lines). In the References Cited section Hoser lists 154 refer- ences (three-and-a-half pages), giving the impression of a well-conducted, literature-based study. However, none of the references is cited in the running text (other than a lengthy list of general references as part of the Introduc- tion), 78 of the references do not refer to Uromastyx tax- onomy or distribution (several are concerned with Leiolepis, others with maintenance of Uromastyx), and 48 references do not refer to Uromastyx at all (including de- scription of statistical methods, herpetofaunal lists outside the distribution of Uromastyx). The only references Hoser appears to actually use are those by Boulenger (1885) and Wilms et al. (2009), from which many lines are copied ver- batim without appropriate attribution. Summary Section D The taxonomic basis for Hoser’s proposals on Uromastyci- nae is a representation of nodes taken fron the phyloge- ny published by Pyron et al. (2013). In total, Hoser’s paper on Uromastyx contains an esti- mated 1490 lines, of which 490 lines are referenced pub- lications and 1000 lines are text (inclusive of title and ab- stract). His diagnoses contribute 556 lines, of which 326 Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK 134 Wolfgang Denzer et al. lines are diagnostic characters. With respect to the latter, of all diagnostic characters mentioned, 255 lines (78%) were identically phrased or copied from previously pub- lished research papers or reviews, with Wilms et al. (2009) and Boulenger (1885) being the main sources. SUMMARY & CONCLUSIONS We analyzed four of Hoser’s publications on agamid lizards and found in all cases significant amounts of copied or plagiarized text to present the diagnostic characters (83% for Draconinae, 82% for Amphibolurinae, 77% for Laudakia, 78% for Uromastycinae). There is no harm per se in repeating diagnostic characters from the older liter- ature and using them in order to define a genus. A species of Draco has a certain number of elongated ribs in the patagium and the nostrils are directed sidewards or up- wards. Similarly, species of Gonocephalus have a gular fold and Chlamydosaurus kingi possess a frill. There ex- ist only a limited number of different expressions to pres- ent certain character sets. However, we think that Hoser’s approach is on a different level that most scientists and editors would consider a sort of plagiarism. We found paragraphs that clearly show that Hoser’s presentation of the diagnosis is a result of a copy-and-paste procedure with typographical errors in exactly the same place in his text where they occurred in the original publication. Further- more, the direct uses of statements from the older litera- ture lack attribution; merely including titles in a bibliog- raphy is not attribution. Meristic characters or statistical values tend to be given with the identical numbers of a source paper (for examples, see the sections on Draco or Uromastyx) although it is clear that Hoser neither took any measurements nor conducted a statistical analysis, as he would not have had access to the same specimens (or any specimens for that matter). By pure repetition of character sets, which are often as old as 125 years, several of Hoser’s diagnoses are rendered inaccurate, inconsistent, or even false. Often a diagnosis consists of more than one character set taken verbatim from two different publications; sometimes as much as half a page is copied in full, or long descriptions are tak- en directly from a previous publication. In at least three cases {Laudakia, Paralaudakia lehmanni, Hypsilurus), sets of characters that were copied by Hoser had been pub- lished in an identical or near-identical manner before, but the original sources are not cited at all! We were able to identify the sources of most (~ 90%) of the diagnostic characters used by Hoser (2012a, 2013, 2014b, c). If the percentage of word-for-word copying of Hoser’s diagnostic characters section is evaluated, this amounts to approximately 80% of his presentation. Even if the whole diagnoses are taken into account for which Hoser typically uses long sentences that have nothing to do with the actual definition of the taxon, the percentage still stands at over 60%. With respect to Hoser’s full pub- lications considered here, approximately 20% {Laudakia paper 11%) of the text constitutes a verbatim copy from other sources. In several cases copied sections exceed 100 lines of identical text, and often full descriptions of species or excerpts from publications concerned with the phyloge- ny or taxonomy of agamid lizards are repeated word-for- word. Hoser’s papers often contain an exhaustive bibliogra- phy which gives the impression of a properly performed literature search. However, we found that actually fewer than 50 publications (out of several hundred referenced) were used. Three publications used are not referenced at all, and none of the publications by his fellow Australian Jane Melville (authored or co-authored) was cited or used, although these contain phylogenies of Laudakia and Di- poriphora as well as nomenclatural proposals preceding those of Hoser. The use of unreferenced material is a clear breach of commonly accepted editorial standards and should be avoided by all means. Hoser’s papers should not have passed any peer review based on the amount of copied text and in our opinion his work constitutes in sev- eral cases a form of plagiarism. Analysing Hoser’s proposed nomenclature we can iden- tify cases where a name is preoccupied and unavailable, where a name is being resurrected that was resurrected be- fore, where names are used that had been very recently synonymized with other genera but for which the litera- ture was overlooked or disregarded. In one case Hoser as- signs a name to a genus that does not include the type species which he places into another genus. In other cas- es, he produces nomina nuda or resurrects a nomen obli- tum incorrectly. He further restricts a type locality with- out identifying a type specimen from that area and selects a holotype for a newly described species that has a bifur- cated tail without any mentioning of this particular fea- ture. With respect to taxonomy, in each of the four papers we find examples of wrong diagnoses, falsely attributed species, and misinterpretation of previously published tax- onomic studies. Furthermore, Hoser omitted several gen- era in his classification schemes (e.g. Harpesaurus, Thau- matorhynchus, Psammophilus and Tympanocryptis) as well as many species (e.g. species of Phrynocephalus and Lophognathus). Presented as they are, Hoser’s taxonom- ic schemes for the subfamilies Amphibolurinae and Dra- coninae, as well as his division of the genera Laudakia and Uromastyx, just constitute a grouping and naming exer- cise within the confines of a particular published phytoge- ny he chooses to follow. The slightest changes in these phylogenies will render them false, in particular as mo- lecular data have not been used to study all genera and species under consideration. Hence, Hoser’s taxonomies and nomenclature acts are highly unstable and little help- Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK Critical review of Hoser’s publications on agamid lizards 135 ful for species assigment. A herpetologist trying to assign a newly collected specimen or an existing museum vouch- er to a specific taxon will still have to look for the origi- nal or subsequent publications where keys are available, and would have to revert to Hoser’s paper(s) only to de- termine his proposed nomenclature - if those names were available. Kaiser et al. (2013) and Kaiser (2013) suggest- ed a suppression of all Hoser names to prevent such case and produced a list with recommended appropriate names. With respect to the taxa dealt with in the present paper we propose to suppress Hoser’s names completely and recommend the usage of generally accepted names which can be found in the Reptile Database (Uetz & Hosek 2015). While available names have to be used according to the Code, a taxonomy does not necessarily have to be accept- ed. If we were to work in the same way as Hoser does we could claim here that Gonocephalus mjobergi and Ptyc- tolaemus share a common character, namely longitudinal gular folds. We could further claim to consider this a synapomorphy not shared by the Japalura variegata / Ori- otiaris group, propose a new tribe excluding Japalura / Oriotiaris with Ptyctolaemus gularis (Peters, 1 864) as the type species and name it accordingly. While our tribe would have a common character to define it, two of Hoser’s tribe diagnoses, which are defined by their con- tent rather than common characters, would become in- valid. Equally we could claim that Hypsicalotes kinabalu- ensis (de Grijs, 1937) has a unique set of characters (which it has) that distinguishes the genus from all other Dracon- inae and remove it from Hoser’s tribe, only to name a new tribe. Although this paper is mainly meant to analyze Hoser’s taxonomy and nomenclature we have to address some is- sues with regard to the Code. There is no requirement for a publication to be peer-reviewed or to comply with any other commonly accepted editorial standards. The ethics recommended by the Code do not have to be adhered to. Even a photograph and short description of the coloura- tion followed by a new name published in a daily news- paper would qualify as valid and therefore the name would be considered available. Zoobank is the official registry of the ICZN. Everyone can register with Zoobank (a vi- able approach and we hope it will stay like this) and sub- sequently register nomenclatural acts. However, Zoobank is not curated and there is no review process in place to check the correctness of submitted data. This literally in- vites pure naming exercises by “harvesting” nodes (nomenclatural vandalism) from a previously published phylogenetic tree. At the end of November 2015, Hoser had 873 nomenclatural acts registered with Zoobank, which on the face of it leads to two different nomencla- tures for many reptilian taxa as his names are not accept- ed by the overwhelming majority of the herpetological community. The ICZN needs to implement provisions to prevent un- scientific and unethical publication of nomenclatural pro- posals to become available. We are convinced that Hoser is abusing the system. The preceding examples provide sufficient evidence to demonstrate his abuses. We strong- ly recommend that the ICZN uses their plenary power to suppress all of Hoser’s nomenclatural acts published in the Australasian Journal of Herpetology. We feel that, if this step is not taken, a large part of the herpetological com- munity will - with great respect for the ICZN and with great regret - continue to use the alternative nomenclat- ural system of the Reptile Database as a reference for available names. CONFLICT OF INTEREST All four authors have submitted taxonomic papers on agamid lizards including nomenclatural changes without acknowledging Hoser’s nomenclature or referencing his publications. Our own publications have been extensive- ly used verbatim by Hoser without asking for permission to do so. The authors therefore have a personal interest to put this on public record. Acknowledgements. The present publication benefited from valuable input provided by colleagues in the field of herpetol- ogy. We are grateful to George Zug and Hinrich Kaiser for a pre- review of the manuscript and for their comments, corrections and suggestions. BIBLIOGRAPHY Amey AP, Couper PJ, Shea GM (2012) Intellagama lesueurii (Gray, 1831), the correct binomial combination for the Aus- tralian Eastern Water Dragon (Sauria, Agamidae). Zootaxa 3390; 65-67 Ananjeva NB, Peters G, Macey JR, Papenfuss TJ (1990) Stel- lio sacra (Smith 1935) - a distinct species of Asiatic Rock Agamid from Tibet. 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Fauna of Arabia 23: 435-468 Wilms TM, Bohme W, Wagner P, Eutzmann N, Schmitz A (2009 “2007”) On the Phytogeny and Taxonomy of the Genus Uro- mastyx Merrem, 1820 (Reptilia: Squamata: Agamidae: Uro- mastycinae) - Resurrection of the Genus Saara Gray, 1845. Bonner zoologische Beitrage 56: 55-99 Zug GR, Brown HH, Schulte II JA, Vindum JV (2006) System- atics of the Garden Eizards, CaJotes versicolor Group (Rep- tilia, Squamata, Agamidae), in Myanmar: Central Dry Zone Populations. Proceedings of the California Academy of Sci- ences, Ser. 4, 57: 35-68 Bonn zoological Bulletin 64 (2): 117-138 ©ZFMK 139 Book Review Lanza B, Funaioli U & Riccucci M (2015) The Bats of Somalia and Neighbouring Areas. Frankfurt Contributions to Natural History 60, Edition Chimaira, Frankfurt/M., 566 pp. ISBN 978-3-89973-447-8. Price: € 78,-. In this voluminous book the authors cover every aspect of the 79 bat species they found associated with Somalia and adjacent areas. There are introductory chapters to the order Chiroptera (with key to the two suborders), subor- der Megachiroptera, followed by suborder Microchi- roptera with a key to the families, then family with key to the genera, and genus with keys to the species, before each species is discussed individually. Each of these chap- ters follows a similar structure with “derivatio nominis” (name derivations and etymology), “composition and dis- tribution”, and “distinctive characters”. Family and genus chapters add paragraphs on “type genus [species]”, “iconography” (= a listing of the figures associated with that chapter), “synonymy” and “common names”, “tax- onomy”, “echolocation calls,” where available, “biology”, “Somali records” and “Personal observations.” Within this pattern the authors provide a great amount of detail on the species, all illustrated with a color drawing of the bat with one wing outstretched (sometimes replaced by color pho- tographs of mounted specimens), up to three (!) sets of pen-and-ink skull drawings per species, nicely showing individual variation, and variously photographs or draw- ings of penis, bacula, palatal ridges, or details of denti- tion, skull, tragi or noseleaves depending on the family’s special characteristics. Quite a bit of information is con- tained in the name derivations, from the Greek and Eat- in roots to interesting details on naturalists or hunter-col- lectors the bat may be named after, and - where available - the bat’s biology. Common names listed include Eng- lish and Italian names but no local Somali names. Well- organized tables identify many of the individual measure- ments of the 3650 museum specimens examined, all giv- en with museum acronyms and numbers, and summary statistics. At the end of the book, under “Addenda,” two species, Kerivoula smithii and Laephotis wintoni are added as “oc- curring in areas nearby Somalia”, followed by 32 detailed distribution maps, some of which covering more than one species, with numbered point localities, all of which iden- tified by name (!) in separate legends. This is then fol- lowed by the Acknowledgements, a detailed six-page Gazetteer with many useful spelling variants and 16 pages of References including literature as recent as some of the species entries from Happold and Happold’s 2013 bat vol- ume in the new Mammals of Africa series, which is great- ly complemented by this new book. As if this wasn’t enough, the book concludes with a sep- arate section by Funaioli and Eanza entitled “An outline of the geography of Somalia” that provides the smaller dis- tribution maps in the main section with much more detail Benedetto [.anza Ugo Funaioli Marco Riccucci The Bats of Somalia and Neighbouring Areas regarding political subdivisions, average yearly rains, and a “Sketch map of the vegetation [zones] of the Somali Democratic Republic,” the latter nicely illustrated with one black & white and 17 color photographs and some more general information on flora and fauna of Somalia. A sep- arate reference section concludes this chapter, which is es- pecially valuable, as it is next to impossible to travel safe- ly in Somalia at this time. If there was ever a modem “Rolls Royce”- equivalent of a “Bats of ...’’-book, with all the possible bells and whis- tles, this must be it. Yet, the publisher. Edition Chimaira, somehow managed to accommodate every lavish aspect of this book and seemingly all special wishes of the au- thors in a still portable size, printed on durable glossy pa- per, which does justice to all the different types of illus- trations, but keeps the price for this rather specialized books within reasonable limits. May this volume find its way to all those interested in Somalia or the fauna of the Horn of Africa in general and bats in particular, and may it serve as an exemplarily illustrated and detailed account for this part of Africa, both areas where the bat-volume of Kingdon’s Mammals of Africa falls short. Jan Decher, Section of Mammalogy, ZFMK Bonn zoological Bulletin 64 (2); 139 ©ZFMK BLANK PAGE