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Revision der Scorpione. II. Scorpionidae und Bothriuridae
... However, no specimen is clearly designated as the holotype in the original description. For reasons given above (see paragraph on Bothriurus burmeisteri Kraepelin, 1894), the male must be considered as lectotype and the female is a paralectotype. ...
... Remarks. Kraepelin (1894) mentioned an additional specimen from the Berlin Museum in the type material. According to the ZMB online catalogue, a paratype from Majunga is present in their collection (ZMB/ Arach-3823). ...
... Remarks. Kraepelin (1894) reported eight specimens in the type series: seven from Reddersburg in the ZMH collections and one from Cape Town in the Natural History Museum of Denmark, Kopenhagen (ZMUC). Prendini (2001) reported seven syntype specimens and designated one female lectotype (ZMB/Arach-7186) in the ZMB, four females paralectotypes in the ZMH (the specimens listed here), one female paralectotype in the BMNH, and one female paralectotype in the ZMUC. ...
Scorpions have always inspired fear and fascination because of the potency of their venoms. Although this ancient arachnid group is relatively small (ca. 2400 species) and has been continuously studied for the past century, the taxonomy is still in a state of flux and the correct identification of species often remains difficult. With more than 725 species and 9000 specimens, the Zoological Museum in Hamburg (ZMH) holds one of the largest and most significant scorpion collections in the world. This collection also contains many historical types described by Karl Kraepelin in the early 20 th century. In order to contribute to a more stable scorpion taxonomy and to assist future scorpion researchers, we present an illustrated and annotated catalogue of the ZMH scorpion collections. The type specimens of 89 species belonging to 10 families are documented, imaged and assessed alongside their primary data. For practical reasons, only the taxa belonging to the parvorder Iurida Soleglad et Fet, 2003 are presented here whilst the Parvorder Buthida Soleglad et Fet, 2003 will be catalogued in a second publication.
... The present study of new samples of hormurid scorpions of the genus Opisthacanthus, subgenus Monodopisthacanthus from Madagascar, has resulted in the discovery of one new species. These were collected from the extreme South-East of the Island, in the Lavasoa humid forest, and are related to Opisthacanthus madagascariensis Kraepelin, 1894 which is known from the western portion of the island. Neverthless, O. madagascariensis is exclusively found in spiny forest thickets and savannah-like formations whereas the new species was found in the humid forest of Lavasoa. ...
... Some comments on the known geographic distribution, the biogeography and ecology of the Malagasy species of Opisthacanthus are included in this paper. Kraepelin, 1894. As already outlined previously (Lourenço & Goodman, 2006) Opisthacanthus madagascariensis was described by Kraepelin (1894) from Majunga (Mahajanga). ...
... Kraepelin, 1894. As already outlined previously (Lourenço & Goodman, 2006) Opisthacanthus madagascariensis was described by Kraepelin (1894) from Majunga (Mahajanga). Subsequently, a second species, O. punctulatus, was described by Pocock (1896) from south central Madagascar. ...
A new species, Opisthacanthus lavasoa sp. n., is described from the Lavasoa Forest, in southeastern Madagascar. The new species shows affinities with both Opisthacanthus madagascariensis Kraepelin, 1894, known from the western portion of the island , and Opisthacanthus ambanja Lourenço, 2014, only known from the extreme north of the island. The new species and O. madagascariensis have similar external morphologies, whereas with O. ambanja the new species shares a similar morphology of the hemispermatophores. Moreover, O. madagascariensis is exclusively found in spiny forest thickets and savannah-like formations , whereas the new species was found in a humid forest. The total number of species in Madagascar is now raised to eleven. Una especie nueva de Opisthacanthus Peters, 1861 (Scorpiones: Hormuridae) del bosque de Lavasoa, sudeste de Madagascar Resumen: Se describe una especie nueva, Opisthacanthus lavasoa sp. n., del bosque de Lavasoa, situado en el sureste de Ma-dagascar. La especie nueva muestra afinidades tanto con Opisthacanthus madagascariensis Kraepelin, 1894, conocido del oeste de la isla, como con Opisthacanthus ambanja Lourenço, 2014, conocido sólo del extremo norte de la isla. La especie nueva tiene una morfología externa similar a la de O. madagascariensis y unos hemispermatóforos semejantes a los de O. ambanja. Por otra parte, O. madagascariensis está presente exclusivamente en manchas de bosque espinoso y formaciones de tipo sabana, mien-tras que la especie nueva se ha encontrado en un bosque húmedo. El número total de especies de Madagascar se eleva así a once.
... Unfortunately, comparaequal, successive, or implied weights and in the additive tively little progress has been made in our understandor nonadditive treatment of multistate characters. The ing of scorpion higher classification since the early preferred hypothesis, a single most parsimonious tree works of Peters (1861), Thorell (1876Thorell ( , 1877, Karsch (1879a,b), Simon (1879Simon ( , 1880, Pocock (1893a), Laurie obtained by analysis with equal weights and 13 ordered (1896a,b), Kraepelin (1891Kraepelin ( , 1894Kraepelin ( , 1899Kraepelin ( , 1905, and Birula Current Classification of the Superfamily Scorpionoidea (1917a,b). Indeed, the suprageneric classification of Re-Latreille 1802 cent scorpions has reached a state in which familial Classification based on Lourenço (1989) and Sissom (1990) assignment of specimens may require prior identifica-Family Bothriuridae Simon 1880 tion to genus (Stockwell, 1989(Stockwell, , 1992. ...
... The (Heteroscorpion ϩ Urodacus) clade was not ob-by character 110 (Appendix 4) and corresponds to the traditional family Scorpionidae (Kraepelin, 1894), be-tained in any of the analyses with implied weights (Figs. 3c-3f), all of which supported Stockwell's (1989) fore the subfamilies Ischnurinae and Heteroscorpioninae were elevated to familial status by Lourenço (1989, placement of Heteroscorpion as sister group of the (Hemiscorpiinae ϩ Ischnuridae) clade, on the basis of charac-1996a). ...
... 13-14. Phoniocercus Pocock 1893: Two species are in this analysis: Centromachetes obscurus Mello-Leitaõ 1932 and Centromachetes pocockii (Kraepelin 1894). The recognised in this bothriurid genus (Sissom, 1990;Lowe and Fet, 2000), both of which were represented second of these is the type species ofCentromachetes. ...
S. A. Stockwell (1989, “Revision of the Phylogeny and Higher Classification of Scorpions (Chelicerata).” Univ. of California, Berkeley) proposed a cladogram and revised classification for the superfamily Scorpionoidea Latreille 1802 (comprising the families Bothriuridae, Diplocentridae, Heteroscorpionidae, Ischnuridae, and Scorpionidae), based on 47 morphological characters and 35 supraspecific terminal taxa, representing genera whose monophyly was implicitly assumed. Given the widespread practice of defining scorpion genera on the basis of plesiomorphic character states, the assumption of monophyly implicit in supraspecific terminal taxa reduces confidence in Stockwell’s cladistic findings and, consequently, his revised suprageneric classification. A re-investigation of scorpionoid phylogeny is presented here, based on 115 morphological characters (including the characters used by Stockwell) and 71 exemplar species. The criterion of “maximal morphological diversity” was employed for exemplar selection. This approach provides a stronger test of monophyly than random exemplar selection. Sixteen cladistic analyses were performed on the scorpionoid data matrix, which varied in the use of equal, successive, or implied weights and in the additive or nonadditive treatment of multistate characters. The preferred hypothesis, a single most parsimonious tree obtained by analysis with equal weights and 13 ordered multistate characters, yielded the scheme of relationships: (Bothriuridae ((Heteroscorpionidae Urodacinae) ((Hemiscorpiinae Ischnuridae) (Diplocentridae Scorpioninae)))). On the basis of these results, revisions are proposed to the existing suprageneric classification of the Scorpionoidea, including new diagnoses, new descriptions, and an illustrated key to the families and subfamilies. Familial status is provided for the scorpionid subfamilies Hemiscorpiinae and Urodacinae.
... Instead, the map includes only specimens about whose correct determination and provenience I am convinced. Among the most dubious published localities are those of Pandinus (Pandinurus) exitialis (Pocock, 1888), P. magrettii Borelli, 1901 and P. pallidus (Kraepelin, 1894). Whereas P. exitialis and P. magrettii definitely are valid species or perhaps even complexes of several species, the validity / taxonomical position of P. pallidus (Kraepelin, 1894) is questionable and invites the possibility that the types are juveniles of another species. ...
... This includes Egypt where the presence of the genus Pandinus has not been subsequently documented (Kraepelin, 1901: 270), and also Kenya and South Sudan (Fet, 2000: 472). The type locality of P. arabicus (Kraepelin, 1894) is "Homran (Arabien, Yemen)" (now Amran, 15°39'45"N 43°56'39"E, city and a province in northern Yemen). However, Vachon & Kinzelbach (1987: 100) did not mention Yemen and stated without any explanation that this species comes from Saudi Arabia. ...
... Fueron descriptas en Cereophonius, o atribuidas a este género, un total de ocho especies. Dos de ellas, e. braehyeentrus Thorell y e. glasioui Bertkau, ambas de Sudamérica, han sido oportunamente transferidas a otros géneros: Urophonius Pocock (Kraepelin, 1894) Y Thestylus Simon (Simon, 1880), respectivamente. Otra especie neotropical, la controvertida Seorpio ehilensis Molina, fue citada por Karsch (1879b) en combinación con Cereophonius, pero hoyes ubicada en el género Bothriurus Peters (Kraepelin, 1894). ...
... Dos de ellas, e. braehyeentrus Thorell y e. glasioui Bertkau, ambas de Sudamérica, han sido oportunamente transferidas a otros géneros: Urophonius Pocock (Kraepelin, 1894) Y Thestylus Simon (Simon, 1880), respectivamente. Otra especie neotropical, la controvertida Seorpio ehilensis Molina, fue citada por Karsch (1879b) en combinación con Cereophonius, pero hoyes ubicada en el género Bothriurus Peters (Kraepelin, 1894). Las cinco especies restantes, todas australianas, responden al concepto genérico de Cereophonius, y eran generalmente aceptadas como válidas: e. squama (Gervais), e. miehaelseni Kraepelin, e. granulosus Kraepelin, e. suleatus Kraepelin y e. kershawi Glauert. ...
The genus Cercophonius Peters is revised, and generic and specific diagnosis are updated. By means of the study of specimens from different localities and the type material, four species (C. michaelseni Kraepelin, C. sulcatus Kraepelin, C. granulosus Kraepelin and C. kershawi Glauert), formerly synonymyzed with the type species C. squama (Gervais), are hereby reinstated as valid; a sixth, new species, C. queenslandae is described. The diagnostic characters used at specific level are: development of ventral carinae in caudal segments I to IV and sternite V; morphology of the hemispermatophore, and pigmentary patterns of tergites and metasoma; other characters, such as the number of pectinal teeth, total lenght and the shape of telson show some differences useful to the determination, although not permiting a clear separation of the species. A key for the species, as well as a discussion on the taxonomic relationships of Cercophonius with Urophonius Pocock and Phoniocercus Pocock (its two closest relatives) are added. A map of the known localities of Cercophonius is also included.
... Early authors (Karsch, 1879;Marx, 1890;Kraepelin, 1894) included in this genus also "Uroctonus phaeodactylus" which now is the type species of Anuroctonus (this, in fact, was a curious recognition of similarity between two uroctonine genera, later obscured by placing both in Vaejovidae and description of more vaejovid genera). Karsch (1879) also included in Uroctonus the third American species, Uroctonus privus, which is now a synonym of Nullibrotheas allenii (Chactidae) (Hjelle, 1972). ...
... Pocock (1893) established a monotypic genus Anuroctonus; the same was done independently by Thorell (1893) who established genus Oncocentrus. Kraepelin (1894) placed "U. phaeodactylus" in Uroctonus, but later (Kraepelin, 1899) accepted Pocock's genus. ...
The systematics of scorpion subfamily Uroctoninae (Scorpiones: Chactidae) is de- scribed and illustrated in detail. Diagnoses of both genera, Uroctonus and Anurocto- nus, and all species and subspecies are provided including an illustrated key and distribution map. For genus Anuroctonus, a comprehensive set of collections was examined, covering its entire geographical range in the USA (California, Nevada, Utah, Idaho) and Mexico (Baja California Norte). A new species, Anuroctonus pococki sp. nov. and its subspecies, A. pococki bajae ssp. nov., distributed in southern coastal California and Baja California Norte, Mexico, are described. It is established that the type species, A. phaiodactylus, is restricted to the Great Basin area of the United States, located in south-eastern California, Nevada, Utah, and Idaho.
... ., ratios 2, 4, 5, 6 separate adult males of N. hierichonticus from those of N. yemenensis), and paired comparisons on th e lower-left half of the matrix lead to the separation of females . : Simon 1878:399, 1880a:29, nee 1902:254, 1910:81, Karsch 1879a:22, Pocock 1903:214, nec Whittick 1941:44, Shulov and Amitai 1958:351, Rosin 1964:177, 1969a:225 , 1969b:71, 1969c:75, 1972:246, 1973:107, Nitzan and Shulov 1966:17, Vachon 1966a:766 , 1966b:214 (in part), 1974:915, 1976:7, nec Vachon 1977:211, Schimdt 1975:2899 : Kraepelin 1894:14 (in part), 1899:98 (in part), nec 1901:270, Borell i 1915:462 (in part ? ), Schenkel 1932:381, Werner 1935a:275 (in part), 1935b:211, Bodenheimer 1937:235, nec Finnegan 1932:92, nec Roewer 1943:224, Bucherl 1960:269, Shulov et al . ...
... Nebo flavipes Simon 1883:249, nec Pocock 1896a:295, 1896b:316, 1903a:214 (in part), Kraepeli n 1899:98, nec 1901:270, Werner 1935b:211, nec Bodenheimer 1937:235, Vachon 1940 :250 (i n part ?), 1965 :308 . Nebo hierichonticus (in part) : Kraepelin 1894:14, Werner 1935a:275, Whittick 1941:44, Roewer 1943:224, Rosin and Shulov 1963:547, Vachon, 1966b:214, Perez 1974 Diagnosis .-Adult females 90-100 mm long (Table 4), adult males unknown . ...
... Only one species, Chaerilus birmanicus (Thorell, 1889), originally described in the genus Chelomachus Thorell, 1889, was clearly reported from Myanmar, originally from Rangoon (Yangôn), Burma. The genus Chelomachus was subsequently synonymised by Kraepelin (1894). ...
... Originally, this species was described as Centrurus phaiodactylus by Wood (1863) from the "Utah Territory" USA, and later transferred to Anuroctonus and placed in the subfamily Iurini of family Iuridae (Pocock, 1893). Later, this genus was transferred to subfamily Vejovini of family Scorpionidae by Kraepelin (1894), to the subfamily Uroctoninae within Vaejovidae (B€ ucherl, 1971), to the subfamily Hadrurinae within Vaejovidae (Stahnke, 1974), and as a member of the superfamily Chactoidea (Francke and Soleglad, 1981). In more recent years, Anuroctonus was a member of Uroctoninae within family Chactidae (Soleglad and Fet, 2004), and lastly, as member of family Iuridae (Prendini and Wheeler, 2005). ...
Scorpions are ancient and historically renowned for their potent venom. Traditionally, the systematics of this group of arthro-pods was supported by morphological characters, until recent phylogenomic analyses (using RNAseq data) revealed most of the higher-level taxa to be non-monophyletic. While these phylogenomic hypotheses are stable for almost all lineages, some nodes have been hard to resolve due to minimal taxonomic sampling (e.g. family Chactidae). In the same line, it has been shown that some nodes in the Arachnid Tree of Life show disagreement between hypotheses generated using transcritptomes and other genomic sources such as the ultraconserved elements (UCEs). Here, we compared the phylogenetic signal of transcriptomes vs. UCEs by retrieving UCEs from new and previously published scorpion transcriptomes and genomes, and reconstructed phyloge-nies using both datasets independently. We reexamined the monophyly and phylogenetic placement of Chactidae, sampling an additional chactid species using both datasets. Our results showed that both sets of genome-scale datasets recovered highly similar topologies, with Chactidae rendered paraphyletic owing to the placement of Nullibrotheas allenii. As a first step toward redressing the systematics of Chactidae, we establish the family Anuroctonidae (new family) to accommodate the genus Anuroctonus.
... Certains de nos prédécesseurs (Kraepelin, 1913) avaient déjà souligné ce fait et utilisé le nombre des granules accessoires de la série basale pour classer certaines espèces de Lychas. ...
... Antecedentes Históricos: Desarrollo y Evolución del DSM Durante el siglo XIX el desarrollo de la nosología psiquiátrica fue restringido intelectual y científicamente, la patología era definida por observaciones de síntomas externos y prevalecían las más antiguas clasificaciones como manía, monomanía, demencia e idiotez (Grob, 1991). No fue hasta finales del siglo XIX que el interés por la nosología resurgió, específicamente por los aportes de Kraepelin (1894), que, aunque no fueron inmediatamente acogidos, contribuyeron al cambio de la visión de unicidad de las circunstancias de las enfermedades por una más generalizada y de probable carácter universal. El primer intento de tener un sistema estandarizado de nosología psiquiátrica surgió en 1908, cuando la Oficina del Censo de los Estados Unidos pidió a la Asociación Americana Médico-Psicológica que creara un Comité de Clasificación de Enfermedades para facilitar la recolección de datos. ...
The text revision of the fifth edition of the Diagnostic and Statistical Manual of Mental Disorders (DSM-5-TR) of the American Psychiatric Association (APA) was published in March 2022. The publication of each version of the DSM generates immediate reactions that increase curiosity, open new (and old) debates and promote discussion among different sectors. In this article we present a summary of the main historical background of the DSM, as well as the criticisms and reflections that have arisen around the publication of each manual. In addition, we present the main changes included in the DSM-5-TR compared to the DSM-5. Lastly, we make final remarks on whether the publication of a new version of the DSM was necessary or not.
... The species is therefore removed from the Saudi Arabian list pending new evidence of its occurrence. (Kraepelin, 1894) This species was described under the protonym Scorpio arabicus by Kraepelin in 1894. It was placed in the genus Pandinus by Kraepelin in 1893, in the genus Pandinurus by Rossi (2015), then in the genus Pandiborellius by Kovařík et al., (2017). ...
A checklist of the scorpion fauna of Saudi Arabia was reviewed and scrutinized published scorpion records considering new taxonomic findings and geographical distribution data. A new checklist to the scorpion fauna of Saudi Arabia is presented. The listed species consisting of 26 species belong to 15 genera and the four families Buthidae (12 genera, 21 species), Diplocentridae (one genus, one species), Hemiscorpiidae (one genus, one species), and Scorpionidae (one genus, three species). The list is dominated by members of the family Buthidae C. L. Koch, 1837 (85.3% of total species) and the genus Compsobuthus is the most species-rich taxon. In addition, a total of 10 dubious records were removed from the list and listed separately pending validations by specimens collected in the country. A dichotomic identification key to the Saudi Arabia scorpion species has been provided.
... Studies on the Malagasy scorpionfauna began in the 19 th century and the first contribution was published by Gervais (1844). Several other isolated publications followed with descriptions of new species and genera (Pocock, 1889(Pocock, , 1890(Pocock, , 1893Kraepelin, 1894Kraepelin, , 1896Kraepelin, , 1900. However, the first comprehensive account of the scorpions of Madagascar was the monography published by Fage (1929), in which a full synopsis was presented. ...
The southwestern portion of Madagascar appears to have one of the highest levels of scorpion diversity on the island. In this contribution, the remarkable diversity of the genus Grosphus Simon, 1880 in this region is discussed. A particular attention is aimed to the area of the Cap Sainte Marie where microendemic geographic patterns are observed. These are tentatively explained in the light of some new biogeographic interpretations. A new species is also described from the Cap Sainte Marie and is characterized by a medium body size and a very pale yellow coloration. While this contribution was in preparation, we learned about the publication of a “most controversial revision” of the genus Grosphus by authors who totally ignored a number of taxonomic particularities of this group and worse, who lack any experience on the ecological and biogeographic patterns observed for the Malagasy fauna. This calls for corrections in which their speculative decisions are refuted.
... In face of Thorell's description, I find it impossible to adopt this opinion [Kraepelin's (1894[Kraepelin's ( , 1899) recognition of Vaejovis intrepidus as type species of the genus]. The measurements given show that V. intrepidus is about twice the size of the average V. mexicanus, the length being 84, the cara- pace 11.5, and the tail 52.5 mm. ...
... KraePeliN (1891) first proposed Cen- trurini, subsequently emended to Centrurinae (KraePe- liN 1899), on Centrurus Ehrenberg, 1829. KraePeliN (1894) later synonymized Centrurus with Heterometrus Ehrenberg, 1828 of family Scorpionidae Latreille, 1802. Centruroides Marx, 1891 was meanwhile proposed, not as a replacement name for Centrurus, but for Buthus exilicauda Wood, 1863, a North American species, and another species that was named but not described (Fet & lowe 2000: 98). ...
All New World buthid scorpions except one South American genus, Ananteris Thorell, 1891, comprise a monophyletic group. The monophyly of two subfamilies, Centruroidinae Kraus, 1955 (= Rhopalurusinae Bücherl, 1971) and Tityinae Bücherl, 1971, proposed to accommodate a subset of these genera, has never been tested. The genera accommodated within Centruroidinae are diverse and poorly defined. Prior to the research presented here, Rhopalurus Thorell, 1876 had a disjunct distribution in the Greater Antilles, the Guiana Shield of northern South America, and northeastern Brazil, where Physoctonus Mello-Leitão, 1934 and Troglorhopalurus Lourenço et al., 2004 also occur. The generic distinction between Rhopalurus and Centruroides Marx, 1890, the most speciose genus of Centruroidinae, distributed from the midwestern United States to northern South America, and throughout the Caribbean, was also unclear. Previous studies suggested Centruroides was paraphyletic with respect to Rhopalurus and vice versa. The study presented here, the first rigorous test of the monophyly of Centruroidinae and its component genera, is based on 90 morphological characters and 4,260 aligned base-pairs of DNA sequence from three mitochondrial and two nuclear DNA loci for 102 terminal taxa, representing 24 species in seven ingroup genera, and nine species in three outgroup genera. Molecular and morphological data, analyzed separately and simultaneously, yielded congruent results. Centruroidinae was monophyletic whereas Tityinae was paraphyletic. Centruroides was monophyletic whereas Rhopalurus was paraphyletic, comprising several monophyletic groups congruent with its disjunct distribution. The results of this analysis justify the redefinition of Rhopalurus and Troglorhopalurus, the revalidation of Heteroctenus Pocock, 1893, and the recently created genera Ischnotelson Esposito et al., 2017 and Jaguajir Esposito et al., 2017. The phylogeny indicates that three distinct types of pecten-sternite stridulation organ evolved in Heteroctenus, Jaguajir and Rhopalurus.
... Quellen: a) BrnuLA 1898BrnuLA , 1903BüRELLI 1913 ;BRULLE 1832;CECC0NI 1913;DI CAPORIACC0 1928, 1948ERBER 1866 ;FRIESE und KöNIGSMANN 1962;GRUB ER 1963KINZELBA CH 1970;KüHNELT 1939;LAMPE 1917;1\1:ENOZZI 1941;MILAN 1962;PAVESI 1877;PENTHER 1906;RüEWER 1943;SCHENKEL 1947;VACHON 1947aVACHON , b , C, 1948VACHON , 1951WERNER 1902WERNER , 1927WERNER , 1928WERNER , 1929WERNER , 1934WERNER , 1935WERNER a, b , 1936WERNER , 1937WERNER , 1941SIMON 1885. Beschreibungen bei BRULLE (1832), KRAEPELIN (1894, 1899), v. UBISCH (1922, VACHON (1947). ...
Considering all the species of scorpions of Southern Europe and the adjacent parts of Asia (inclu-ding key for determination and distribution maps) the six species occurring in the countries around the Egean Sea are revised. They belong to the Buthidae (1), Vejovidae (1), and Chactidae (4). Given are synonyms, morphology as far as it is useful for identification, subspecies, ecology, and distribution. lt is suggested that Euscorpius carpathicus auct. bedivided into two species: E. (E.J car-pathicus (L.) s. str., and E. (E.J mesotrichus HADZI. The coincidence of the pattern of distribution of morphological characteristics with the results of the reconstruction of distribution during the Tertiary period encouraged the establishment of a new theory of the phylogeny of the Euscorpiinae and Calchinae. Inhaltsverzeichnis
... The presence of an eyespot was not conclusively established as a diagnostic character for Chaerilidae until now. Kraepelin (1894Kraepelin ( , 1899 and Tikader and Bastawade (1983) referred to an amber or yellow spot near the lateral ocelli in their diagnoses but many other authors ignored the structure (Pocock, 1893(Pocock, , 1900Stockwell, 1989;Sissom, 1990;Kovařík, 2000;Soleglad and Fet, 2003;Kovařík and Ojanguren Affilastro, 2013). Loria and Prendini (2014) defined the lateral ocellus pattern in Chaerilidae as Type 2A, comprising a mediolateral major ocellus (MLMa) and a posterolateral major ocellus (PLMa), but also observed several aberrant conditions including In the present study, an eyespot was observed in all 733 individuals examined, representing 39 (93%) species of Chaerilidae, confirming that it is ubiquitous across the family. ...
... COMMENTS. Pandinus pallidus was based on two juveniles incorrectly cited as adults, a male and a female (Kraepelin, 1894: 60, Kovařík, 2009. Kovařík (2012: 20) wrote that "taxonomical position of P. pallidus is questionable and invites the possibility that the types are juveniles of another species". ...
... De plus, schaumi avait été décrit de l'Inde, localité type sans doute fausse ; cependant, la localité type reste incertaine, mais se trouve proba¬ blement à l'intérieur de la région guyanaise. Plus récemment, Mello-Leitâo (1945) cite Broleochactas schaumi, suivant l'opinion de Kraepelin ;Gonzalez-Sponga (1978) Hadrurochactas schaumi (Karsch, 1880) Cette espèce est de petite taille (voir tabl. I) ; la queue est très large et forte par rapport au corps ( fig. ...
... 396 . Kraepelin "1893" (1894), p . 11 . ...
... However, for the purposes of the present study, it is important to note that species previously placed in the genera Heterometrus or Scorpio, viz. O. boehmi (Kraepelin 1896(Kraepelin , 1899Pocock 1900b) and O. carinatus (Peters 1861(Peters , 1862, were unequivocally placed in Opistophthalmus, following Kraepelin (1894Kraepelin ( , 1913 Fig. 7. A, The optimal tree obtained by simultaneous analysis of the morphological data (Table 3) and aligned molecular data (Accessory Material) under the weighting regime that minimised length and maximised fit. This MPT was obtained by analysis with implied weights under k = 6 and gaps included (Table 10). ...
A cladistic analysis of relationships among the genera of Scorpionidae Latreille, 1802—Heterometrus
Ehrenberg, 1828; Opistophthalmus C. L. Koch, 1837; Pandinus Thorell, 1876; and Scorpio Linnaeus, 1758—based on morphology and DNA sequence data from loci of three genes in the mitochondrial genome (12S ribosomal DNA (rDNA), 16S rDNA and cytochrome oxidase I) and one gene in the nuclear genome (28S rDNA) is presented. The analysis makes use of exemplar species, specifically selected to test the monophyly of the genera, rather than supraspecific terminal taxa. Other methods used in the analysis are justified in the context of a discussion of current methods for phylogenetic reconstruction. Relationships among the scorpionid genera are demonstrated to be as follows: (Opistophthalmus (Scorpio (Heterometrus + Pandinus))). This reconstruction identifies Opistophthalmus as the basal lineage of the Scorpionidae, rather than the sister-group of Scorpio. Revised descriptions, diagnoses and a key to identification of the four scorpionid genera are provided, together with a summary of what is known about their ecology, distribution and conservation status.
... [example from Valparafso, ZMH, misidentified]; Cekalovic 1983 (part.):51 [reference to Kraepelin 1894] ...
Urophonius transandinus, new species belonging in the brachycentrus species-group is described. It can be separated from other species in the group by the pigmentation pattern of mesosoma and ventral side of metasoma, the number of pectinal teeth, and the arrangement of the ventral submedian keels of caudal segment I. Additionally, hemispermatophores of the new species lack a posterodistal fold (present in the rest). Records of U. transandinus sp.nov. come from the Chilean provinces Valparaíso, San Antonio, San Felipe de Aconcagua, Santiago and Cordillera. The probable zoogeographical links of Central Chile and the "peripampasic track" (Argentina – Uruguay – southern Brazil), as suggested by the geonemy of the brachycentrus group, are discussed.
... The scorpion fauna of central Chile is still poorly known, with most of the published literature consisting of short species descriptions dating from the late 19th century and early 20th century (Pocock 1893(Pocock , 1898Kraepelin 1894Kraepelin , 1911Lönnberg 1897;Mello-Leitão 1932;Werner 1939). Little subsequent taxonomic work on this fauna is available and the identity of most of the species from this area needs to be confirmed. ...
Brachistosternus (Leptosternus) aconcagua new species, from the Chilean central Andes, is described. Brachistosternus (L.) chilensis Kraepelin 1911 is redescribed. Both species are related to other Andean species of the genus and with other species from central and northern Chile. RESUMEN. Se describe a Brachistosternus (Leptosternus) aconcagua nueva especie de los Andes centrales Chilenos. Se redescribe a Brachistosternus (L.) chilensis Kraepelin 1911. Ambas especies se encuentran relacionadas con las especies andinas del género y con algunas especies del centro y norte de Chile.
... 396 . Kraepelin "1893" (1894), p . 11 . ...
... 14), and Broteochactas, with eta norma l and et 5 small (positions "b" and "c" in Fig. 14) is based on a false homology an d certainly does not justify the recognition of two separate genera . : Kraepelin 1894:176-177, 1899:173, 1912:53 Pocock 1897:365-366 , 1900Mello-Leitao 1932:32, 1945 : Roewer 1943 :237 ;Waterman 1950 :169 ;Kjellesvig-Waering 1966 :126 . (Fig . ...
The first phylogenetic analysis of genus Urophonius is performed based on morphological characters. Three monophyletic species groups are recovered. Species of the granulatus species group and Urophonius mondacai have summer or spring activity patterns. The exochus and brachycentrus species groups form a monophyletic group with a winter activity pattern, which is so far unique in scorpions from temperate and cold areas. Additionally, two new Patagonian species of Urophonius are described, one from Argentina and one from Chile. Finally, we discuss the southernmost distributions of scorpions in continental Patagonia. We suggest that the southern limit of the group could be related not only with a decrease in temperature, but also with an increase of precipitation.
Euscorpius tauricus (C. L. Koch, 1837) was previously known only from the Crimea Peninsula, Ukraine. We report an unexpected presence of this species in the Cyclades Islands (Greece) and northwestern Anatolia (Turkey). In addition we designate a neotype for this species. We synonymize Euscorpius carpathicus aegaeus Di Caporiacco, 1950 syn. n., from Antiparos Island and Euscorpius rahsenae Yağmur et Tropea, 2013 syn. n., from Anatolia, with E. tauricus. In addition, we describe two new species related to E. tauricus, from the Cyclades Islands: E. curcici sp. n., from Ios and Sikinos Islands, and E. amorgensis sp. n., from Amorgos Island. Identity and level of divergence of these taxa is confirmed by multiple DNA markers.
The new species Brachistosternus paulae is described. This is the southernmost species of the genus, occurring in southern Patagonia in Santa Cruz Province. It can be distinguished from all the other species of the genus by the presence of only 4 ventral trichobothria on the pedipalpal chela, and by the shape of the hemispermatophore, in which the cylindrical apophysis is shorter than the laminar apophysis; all internal spines are absent, except for the row of spines, which in some specimens is vestigial. This species can not be included in any of the known subgenera due to its trichobothrial pattern; nevertheless it is closer to the subgenus Leptosternus on the basis of its remaining morphology. A key for the species of the genus in the country is provided together with some data on them. The hemispermatophores of the three subgenera are compared and two maps detailing the distribution of the species in the country are provided.
A synopsis is presented of the scorpion fauna documented in the Réserve spéciale d'Ankarana in northern Madagascar, an area of limestone karst. Two new species of liochelid scorpions, Opisthacanthus milloti n. sp. and Opisthacanthus pauliani n. sp., are described based on specimens collected in the reserve. One of these, O. pauliani n. sp. may represent the second troglobitic scorpion species found in Madagascar, and the first one belonging to the genus Opisthacanthus Peters, 1961 anywhere in the world. © Publications Scientifiques du Muséum national d'Histoire naturelle.
Two new species of Chaerilus aredescribed from Southeast Asia: C. anneae sp.n. from southern Vietnam and C. kampuchea sp.n. from Cambodia. These new species are compared to the other Chaerilus species known from Southeast Asia and Indonesian islands. This study suggests that different species are not too much distinct morphologically, rather corresponding to micro-endemic populations. Molecular studies, now in preparation, are deemed to bring further evidence to favour or refute this hypothesis.
On Hadogenes angolensis Louren o, 1999 syn. n. (Scorpiones, Liochelidae), with a redescription of H. taeniurus (Thorell, 1876). - The flat rock scorpion, Hadogenes taeniurus (Thorell, 1876), is redescribed, based on an examination of approximately 250 specimens from Angola and Namibia. Hadogenes angolensis Lourenco, 1999 is demonstrated to be a junior synonym. The distribution of H. taeniurus is mapped, and notes on its ecology and conservation status are provided. The synonymy of Hadogenes bifossulatus Roewer, 1943 with Hadogenes tilyrus (Simon, 1888), rather than with H. taeniurus, is confirmed.
Scorpions previously assigned to the genus Liocheles Sundevall, 1883, of the family Hormuridae Laurie, 1896, are widely distributed in the tropical forests of the Indo-Pacific region. Revisionary systematics of these poorly known scorpions has revealed a tremendous diversity of species. As part of an ongoing investigation, the first analysis of Indo-Pacific hormurid scorpion phylogeny based on morphological data scored for all currently recognized species of Hormiops Fage, 1933, Hormurus Thorell, 1876, and Liocheles, is presented. The taxonomy of these scorpions is reassessed and their biogeography reinterpreted in the light of the phylogeny. Phylogenetic, morphological, and distributional data support the revalidation of Hormiops and Hormurus, previously synonymized with Liocheles. The phylogeny indicates that the Australasian hormurids are more closely related to the Afrotropical and Neotropical hormurids than to the Indian hormurids, as previously proposed, refuting the “out-of-India” origin of Asian hormurids. A recent paleogeographical hypothesis, the “Eurogondwana model”, is supported instead. According to this hypothesis, hormurid scorpions colonized Laurasia from Africa via the Apulia microplate (Europa terrane) in the Cretaceous, subsequently colonized the Australo-Papuan archipelago in the early-mid Cenozoic, and then went extinct in the Northern Hemisphere during the second half of the Cenozoic. These results suggest that, contrary to the traditional paradigm, dispersal and extinction may affect spatial and temporal biotic distributions as much as vicariance, even in animals with limited vagility, such as scorpions.
We review the taxonomy of the Hadruroides Pocock, 1893 (Iuridae: Caraboctoninae), scorpions of Peru, describe six new species from the north of the country, and report new records of other poorly known species. The description of these species raises to 16 the number of described species in the genus, 13 of which occur in Peru. Four species inhabit dry forest in northern Peru: H. charcasus (Karsch, 1879); H. chinchaysuyu, n. sp.; H. geckoi, n. sp.; H. leopardus Pocock, 1900. Three species occur in inter-Andean valleys along the Cordillera: H. bustamantei Ochoa and Chaparro, 2008; H. carinatus Pocock, 1900; H. mauryi Francke and Soleglad, 1980. Six species inhabit desert along the Pacific coast: H. aguilari Francke and Soleglad, 1980; H. graceae, n. sp.; H. juanchaparroi, n. sp.; H. lunatus (L. Koch, 1867); H. tishqu, n. sp.; H. vichayitos, n. sp. Most species of Hadruroides have restricted distributions, except H. charcasus and H. lunatus, which are apparently more widely distributed. We consider it necessary to reassess all previous records of the latter two species, because we suspect several are based on misidentifications.
The male of Megacormus
granosus is described for the first time and the female redescribed. A homology scheme proposed recently is applied to hemispermatophore structures. The specimens were collected in an oak forest from Pico de Orizaba Volcano at an average altitude of 2340 m. All adult males were collected by pitfall traps, whereas all adult females and both sex immatures were collected using Berlese funnels, suggesting that males are comparatively more mobile within the leaf litter layer, probably due to mating season.
Scorpions of Djibouti, Eritrea, Ethiopia, and Somalia are revised, with a key and geographic distributions given for all the species. Lectotypes are designated for Buthus minax L. Koch, 1875, Buthus hottentotta tigrinus Caporiacco, 1937, Parabuthus granimanus Pocock, 1895, Parabuthus granimanus fuscicauda Caporiacco, 1947, Parabuthus mixtus obscurior Caporiacco, 1941, Parabuthus mixtus Borelli, 1925, Scorpio cavimanus Pocock, 1888, Pandinus intermedius Borelli, 1919, Scorpio pallidus Kraepelin, 1894, and Scorpio smithi Pocock, 1897. Neobuthus Hirst, 1911 is synonymized with Butheolus Simon, 1882. The following species are synonymized: Neobuthus berberensis Hirst, 1911 with Butheolus ferrugineus Kraepelin, 1898; Buthus hottentotta tigrinus Caporiacco, 1937 with Hottentotta minax (L. Koch, 1875); Buthus fuscitruncus Caporiacco, 1936 with Hottentotta trilineatus (Peters, 1862); Parabuthus granimanus fuscicauda Caporiacco, 1947 with Parabuthus granimanus Pocock, 1895; Parabuthus heterurus stefaninii Caporiacco, 1927 with Parabuthus heterurus Pocock, 1897; Parabuthus leiosoma abyssinicus Pocock, 1901 and Parabuthus leiosoma dmitrievi Birula, 1903 with Parabuthus leiosoma (Ehrenberg, 1828); Parabuthus mixtus Borelli, 1925, Parabuthus mixtus obscurior Caporiacco, 1941, and Parabuthus zavattarii Caporiacco, 1939 with Parabuthus pallidus Pocock, 1895; Uroplectes fischeri caporiaccoi Fet, 1997 and Uroplectes patrizii Caporiacco, 1936 with Uroplectes fischeri (Karsch, 1879); Scorpio gregoryi Pocock, 1896 with Pandinus (Pandinurus) exitialis (Pocock, 1888); Brotheas hirsutus L. Koch, 1875 and Scorpio africanus subtypicus Kraepelin, 1894 with Pandinus (Pandinurus) magrettii Borelli, 1901; and Pandinus intermedius Borelli, 1919 and Pandinus citernii Borelli, 1919 with Pandinus (Pandinus) phillipsii (Pocock, 1896). Uroplectes pardii sp. n. from Somalia and Parabuthus eritreaensis sp. n. and Pandinus (Pandinops) eritreaensis sp. n. from Eritrea are described. Hottentotta minax niloticus Birula, 1928 is elevated to a full-species status. First records are established for Leiurus quinquestriatus Ehrenberg, 1828 in Somalia, Lychas obsti Kraepelin, 1913 in Ethiopia and Somalia, Pandinus (Pandinops) hawkeri Pocock, 1900 in Ethiopia, and Pandinus (Pandinurus) exitialis (Pocock, 1888) in Somalia.
Manuskript eingelangt am 12. Dezember 1991 Zusammenfassung Im Verlauf von arachnologischen Untersuchungen im östlichen Mittelmeerraum wurde den Skorpionen besondere Beachtung geschenkt. So konnten die Arten Mesobuthus gibbosus (BRULLE), lurus dufoureius (BRULLE), Iurus dufoureius asiaticus BIRULA, Euscorpius carpathicus carpathicus (LINNÉ), Euscorpius carpathicus candiota BIRULA, Euscorpius mesotrichus HADÈI, Euscorpius min-grelicus (KESSLER) und Euscorpius (Polytrichobothrius) italicus (HERBST) für dieses Gebiet fest ge-stellt und zahlreiche neue Fundplätze erbracht werden. Neben kurzen Darstellungen über die jeweilige Ökologie und allgemeine Verbreitung wurden Bemerkungen angestellt, die zur Klärung der systematischen Stellung der einzelnen Arten beitragen sollen.
Une nouvelle espèce appartenant au genre Chaerilus Simon, 1877 est décrite d’une grotte située dans le district de Dien Bien, à l’ouest de Dien Bien Phu City, au nord du Vietnam. La nouvelle espèce est morphologiquement similaire à d’autres espèces de Chaerilus distribuées uniquement dans le Sud du Vietnam et du Cambodge. Ceci suggère un cas d’espèces vicariantes entre les populations distribuées dans le Nord et le Sud de la péninsule Sud-Est asiatique.
Scorpions previously assigned to the genus Liocheles Sundevall, 1883, of the family Hormuridae Laurie, 1896, are widely distributed in the tropical forests of the Indo-Pacific region. Revisionary systematics of these poorly known scorpions has revealed a tremendous diversity of species. As part of an ongoing investigation, the first analysis of Indo-Pacific hormurid scorpion phylogeny based on morphological data scored for all currently recognized species of Hormiops Fage, , Hormurus Thorell, 1876, and Liocheles, is presented. The taxonomy of these scorpions is reassessed and their biogeography reinterpreted in the light of the phylogeny. Phylogenetic, morphological, and distributional data support the revalidation of Hormiops and Hormurus, previously synonymized with Liocheles. The phylogeny indicates that the Australasian hormurids are more closely related to the Afrotropical and Neotropical hormurids than to the Indian hormurids, as previously proposed, refuting the “out-of-India” origin of Asian hormurids. A recent paleogeographical hypothesis, the “Eurogondwana model”, is supported instead. According to this hypothesis, hormurid scorpions colonized Laurasia from Africa via the Apulia microplate (Europa terrane) in the Cretaceous, subsequently colonized the Australo-Papuan archipelago in the early-mid Cenozoic, and then went extinct in the Northern Hemisphere during the second half of the Cenozoic. These results suggest that, contrary to the traditional paradigm, dispersal and extinction may affect spatial and temporal biotic distributions as much as vicariance, even in animals with limited vagility, such as scorpions.
abStraCt The scorpion genus Diplocentrus Peters, 1861, comprising more than 50 species, most of which are endemic to Mexico, is the most diverse in the family Diplocentridae karsch, 1880 (Santibáñez-lópez et al., 2011). Hoffmann (1931) divided the Mexican species into two groups, the whitei group and the keyserlingi group, based largely on differences in size and coloration. Francke (1977) redefined these groups. The whitei group, renamed the mexicanus group because it included the type species of the genus, comprised species with short cheliceral fingers and the pedipalp femur wider than high. The keyserlingii group comprised species with long cheliceral fingers and the pedipalp femur higher than wide. Several new species of Diplocentrus were since described, but no attempt was made to synthesize the taxonomy of the species assigned to either group or further clarify the validity of the groups. in the present contribution, the species of Diplocentrus with the pedipalp femur higher than wide are reviewed. an operational diagnosis is provided for the keyserlingii group. Diplocentrus formosus armas and Martín-Frías, 2003, previ-ously synonymized with Diplocentrus tehuano Francke, 1977, is reinstated. revised, updated diag-noses are provided for all previously described species and three new species, Diplocentrus
Examination of an extensive material that includes the type series and recently collected specimens enable a precise reassessment of the status of several taxa belonging to the genus Liocheles Sundevall, 1833 (Scorpiones, Liochelidae). Two species previously in synonymy, i. e. Liocheles boholiensis (Kraepelin, 1914) and Liocheles neocaledonicus (Simon, 1877), are revalidated. Liocheles australasiae longimanus (Werner, 1939) is elevated to species rank and Liocheles australasiae brevidigitatus Werner, 1936 is synonymized with Liocheles australasiae (Fabricius, 1775). L. boholiensis (Kraepelin, 1914), L. neocaledonicus (Simon, 1877) and L. longimanus (Werner, 1939) are thoroughly redescribed, diagnosed and illustrated, and their distribution ranges are accurately mapped.
The family Hemiscorpiidae is closely related to the Liochelidae. Within the Hemiscorpiidae, the Oriental species are particularly interesting. Most of them exhibit highly derived characters in comparison to their African relatives. Males possess a strongly elongated metasoma and a similarly elongated telson bearing a pair of tuberculiform processes at the base of the aculeus. Furthermore, Hemiscorpius lepturus Peters, 1861, which occurs in Iraq and Iran, is known to have an extremely virulent venom with cytotoxic and haemolytic components. It is responsible for severe dermonecrotic scorpionism in southern Iran. This is the only non-buthid scorpion that is potentially lethal. In this paper an overview of the species of Hemiscorpius in Iran is presented with revised diagnoses and descriptions. Two new species from western Iran, H. enischnochela sp. n. and H. acanthocercus sp. n., are described. The genus Habibiella Vachon, 1974 is synonymised with Hemiscorpius Peters, 1861. A thorough analysis of hemispermatophores shows close phylogenetic relationships with several genera of the family Liochelidae. A hypothesis on the geological events that probably triggered the present distribution of Hemiscorpius is finally proposed.
The purpose of this application under Articles 23.9.3 and 81.2.3 of the Code is to conserve the well known specific name of Bothriurus alticola Pocock, 1899 for a scorpion (family BOTHRIURIDAE) from Argentina by giving it precedence over the little used subspecific name Cercophonius braehycentrus bivittatus Thorell, 1877. Pocock's nominal species is the type of Orobothriurus Maury, 1976, a genus currently including 10 species from Andean and sub-Andean localities in Argentina and Peru, most occurring at high altitude.
Several previous nomenclatural corrections are analyzed and the valid name determined for each case. These include (a) emendations and subsequent spellings: Brachistosternus weijenberghi (Thorell, 1877a) (Bothriuridae), Urophonius iheringi Pocock, 1893 (Bothriuridae), Lisposoma jose- hermana Lamoral, 1979 (Bothriuridae), Chactas raymondhansi Francke & Boos, 1986 (Chactidae), Tityus fuhrmanni Kraepelin, 1912 (Buthidae), Parabuthus liosoma (Ehrenberg, 1828) (Buthidae), Hadrurus concolorous Stahnke (Iuridae), Vachoniolus minipectenibus (Levy, Amitai & Shulov, 1973); (b) incorrect reversal of precedence: Hadogenes hahni (Peters, 1862) n. comb. (Lioche- lidae); (c) genus and species names that never were available: Pucha, Puchale, Repucha (Francke, 1985; Fet, 1997), Transbothriurus (Mello-Leitão, 1945; Lowe & Fet, 2000), Scorpio maurus puni- cus Fet, 2000 (replacement name for Scorpio maurus tunetanus Birula, 1910) (Scorpionidae)
The nitidulus group of Vaejovis is defined on the basis of pedipalp chela morphology and dentition , trichobothrial patterns, and carinal structure of the pedipalp chela and the metasoma . Five specie s are treated herein : V. mexicanus decipiens Hoffmann is elevated to specific status and transferred t o the nitidulus group ; V. intermedius Borelli, V. nigrescens Pocock, and V nitidulus C. L. Koch ar e all considered valid species and redescribed ; and V. minckleyi Williams is transferred to the grou p and a revised diagnosis given .
Sperm packages are widespread in the order Scorpiones, but absent in the family Buthidae. The morphology of sperm packages is diverse and apparently has phylogenetic information. The objectives of this work were to show diversity of sperm packages and to provide a quantitative basis for using sperm packages’ morphology as a taxonomic character. For this, we conducted a morphological analysis and comparison of the different sperm packages of species of the family Bothriuridae. The seminal content from males of species of Bothriuridae was studied. Specimens from Iuridae, Buthidae, Euscorpiidae, Liochelidae, Scorpionidae, Vaejovidae, Chaerilidae, and Chactidae were used for comparison. Digital images of sperm packages were measured and statistically analysed based on the following variables: total length, head width, head–body angle, total area, and head length. Pairs of variables were also contrasted, and all the variables were correlated with the current phylogenetic hypothesis for Bothriuridae. High morphological diversity and variability in measures was observed. In general, measurements were similar within each genus, but differed amongst genera. Cane-like sperm packages are very common in species of the family Bothriuridae. Species from Bothriurus show a wide range of sperm package shapes, some of them shared with Timogenes and Vachonia species, supporting the idea of nonmonophyly of the genus. Many species showed sperm package dimorphism inside a single male. Some of the analysed features fit well with the phylogenetic hypothesis in Bothriuridae, and the general package shape shows high correlation with scorpion phylogeny in other families. Bent and round packages are the most common amongst the different families. Sperm packages are not developed in Chaerilidae, as in Buthidae. This is the first morphological and comparative analysis of sperm packages in scorpions, and reveals much greater diversity in this trait than previously known. Our results reinforce the idea that the study of morphology of sperm packages would contribute characters for scorpion phylogeny at different levels.
The endemic Namibian genus Lisposoma Lawrence, 1928, comprising two described species, represents one of two basal African genera in the Gondwanan family Bothriuridae Simon, 1880. All other, more derived genera of Bothriuridae occur in South America, India and Australia. Although the phylogenetic position of Lisposoma has become an increasingly contentious subject, three cladistic analyses based on morphological data have confirmed that it is a basal bothriurid. In view of those findings, the present contribution serves to revise the generic diagnosis of Lisposoma, last revised by Lamoral (1979), who placed the genus in the Scorpionidae Latreille, 1802 and constructed his diagnosis accordingly. A considerable number of new specimens, many representing new records for the two species of Lisposoma, have also accumulated since Lamoral's (1979) revision. These new data justify the provision of revised diagnoses and descriptions for the two species, together with a key to their identification, brief summaries of their ecology and conservation status, and a distribution map plotting all known locality records.
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