Taxonomic revision of modern Doras Lacepède, 1803 (Siluriformes: Doradidae), with descriptions of three new species
Mark H. Sabaj Pérez
Department of Ichthyology, The Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103-1195, USA
José L. O. Birindelli
Museu de Zoologia da Universidade de São Paulo, Caixa Postal 42494, 04218-970, São Paulo, SP, Brazil
ABSTRACT.—Modern Doras is newly diagnosed among Doradidae by the unique combination of maxillary barbels long and fimbriate; mesethmoid with anterior lateral margins converging towards pointed tip; single anterior cranial fontanel contained largely within frontals and anteriorly by mesethmoid; anterior nuchal plate wide, subpentaganol to subhexagonal, sharing broad lateral suture with epioccipital and isolating supraoccipital from middle nuchal plate; nuchal foramina absent; coracoid process short, posterior tip finishing well short of that of postcleithral process; dentaries typically with acicular teeth; and skin beneath postcleithral process perforated with conspicuous pores. One fossil species, †D. dioneae, and two nominal modern species, D. carinatus and D. micropoeus, are recognized as valid with three additional modern species, D. phlyzakion, D. higuchii and D. zuanoni, newly described from the middle Amazon and tributaries, lower Amazon tributaries and Rio Araguaia (Tocantins drainage), respectively. The five modern species are placed into two phyletic groups: the phlyzakion group (D. phlyzakion and D. zuanoni) typical of lowland, lentic habitats and the carinatus group (D. carinatus, D. higuchii and D. micropoeus) typical of upland, lotic habitats. Species of the phlyzakion group are characterized in part by numerous pores in skin on ventral surfaces, a trait unique among doradids and perhaps among all catfishes. The occurrence of D. carinatus in the Orinoco is newly confirmed and suggests a historical link between right-bank tributaries of the lower Orinoco (e.g., Caroní) draining the western Guiana Shield and more eastern rivers (e.g., Cuyuní-Essequibo) draining the Shield directly into the Atlantic Ocean. A key to modern species is provided and a neotype is designated for Silurus carinatus Linnaeus 1766.
Species nova: Doras higuchii, Doras phlyzakion, Doras zuanoni Sabaj Pérez and Birindelli
The taxonomic history of Doras Lacepède is one of repeated expansion and contraction. Lacepède (1803:116) created Doras (his Genus 164) for two Linnean species: Silurus costatus Linnaeus 1758 and Silurus carinatus Linnaeus 1766. Bleeker (1858:53–54) expanded Doras to include 19 valid species (plus one as questionable). In the same work, Bleeker (1858:48) was first to recognize a higher-level taxon, “Phalanx Doradini” (“Subfamilia Callichthyoidei”), for species currently placed in Doradidae and within this group he recognized three genera: Doras, Pseudodoras (4 species) and Hemidoras (1 species). Bleeker (1862:5) later designated Doras carinatus the type species of the genus and subsequently (Bleeker, 1863) restricted Doras to seven species in his early revision of doradids. Eigenmann and Eigenmann (1888:158), evidently overlooking Bleeker’s designation, transferred Doras carinatus to the genus Hemidoras Bleeker and re-expanded Doras to include 24 species in seven subgenera. Later Eigenmann and Eigenmann (1890) similarly recognized 24 species in Doras and designated Silurus costatus Linnaeus the type species of the genus while maintaining Linnaeus’ carinatus in Hemidoras. Miranda Ribeiro (1911) restricted Doras to 14 species and created a new monotypic genus, Mormyrostoma, for the Linnaean carinatus. Eigenmann (1925), in his remarkable monograph on the family Doradidae, recognized Bleeker’s earlier designation of Silurus carinatus as the type species of Doras and restricted the genus to seven species: D. carinatus, D. micropoeus, D. punctatus, D. fimbriatus, D. lipophthalmus, D. microstomus and D. brevis. Eigenmann also noted in a footnote that two species, Doras microstomus and D. brevis, might belong in Trachydoras. Gosline (1945:23) removed D. lipophthalmus and Fernández-Yépez (1968:32) removed D. microstomus to Hassar and Anduzedoras, respectively. Sabaj and Ferraris (2003) further restricted Doras to include but two valid species, Doras carinatus and D. micropoeus Eigenmann 1925, removed D. brevis to Trachydoras, and treated D. punctatus and D. fimbriatus with Oxydoras eigenmanni as valid species incertae sedis in Doradidae. Finally, Sabaj Pérez et al. (2007) described a new fossil species, †Doras dioneae, based on a partial left pectoral girdle with articulated pectoral-fin spine from the Late Miocene (ca. 8 Ma) Urumaco Formation, Falcón State, Venezuela.
Prior to this study, the two modern species of Doras were known to inhabit Atlantic coast drainages from the Essequibo River, Guyana, to Amapá State in northeastern Brazil (Le Bail et al., 2000; Sabaj and Ferraris, 2003; Sabaj Pérez et al., 2007). Examination of museum specimens and recent expeditions to the upper Rio Xingu funded by the All Catfish Species Inventory (http://silurus.acnatsci.org) resulted in the discovery of three new species of Doras from the middle and lower Amazon basin, and Rio Araguaia, Tocantins basin. In this paper we rediagnose and redescribe modern Doras and two modern nominal species; diagnose and describe three new modern species; provide a key to modern species; designate a neotype for Silurus carinatus Linnaeus 1766; define two species groups within modern Doras; and briefly discuss the biogeographical implications of the distribution of the genus.
MATERIALS AND METHODS
Measurements were made to the nearest 0.01 mm using dial calipers; methodology follow Sabaj (2005) and Sabaj Pérez et al. (2007) with the following additions and exceptions: head length = distance from snout tip to dorsalmost point of gill opening; body depth at anal-fin origin = vertical depth from anal-fin origin to dorsal margin of body or adipose fin; caudal peduncle length = horizontal midlateral distance from vertical through base of last anal-fin ray to posterior margin of hypural plates; dorsal spine length = distance from point on midlateral base of spine even with dorsal margin of body to distal bony tip; snout length = from tip of snout to anteriormost margin of eye; opercle width = distance between dorsalmost points of gill openings; nuchal shield width = minimum transverse width of nuchal shield (across middle nuchal plate); maxillary barbel length = from inferior base of barbel (where it meets labial tissue) to distal tip; outer mental barbel length = from base of outer mental barbel to distal tip. Standard length (SL) is expressed in mm; other measurements are expressed as percentages of standard length or head length (e.g., subunits of head).
Midlateral scute counts were taken on the left side of body (when permissible) and begin with infranuchal scute (connected dorsally to posterior nuchal plate and internally to first complete rib, borne on sixth vertebra). The nuchal shield is considered herein to be composed of the anterior, middle, and posterior (paired) nuchal plates (dermal bones) which are superficial expansions of the supraneural, first and second pterygiophores (endochondral bones), respectively. Specimens are designated as alc (alcohol), sk (dry skeleton) and cs (cleared and stained prepared according to Taylor and Van Dyke, 1985) with measurements recorded as SL unless specified as TL. Museum abbreviations follow Ferraris (2007); collection dates for 1908 expedition to Guyana led by Carl H. Eigenmann follow Hardman et al. (2002). See Sabaj Pérez et al. (2007) for doradid comparative material examined. Only modern species of Doras are described and redescribed herein; for complete description of fossil †D. dioneae see Sabaj Pérez et al. (2007). Species description complete for D. carinatus; subsequent species descriptions may refer to D. carinatus when morphology is similar.
Doras Lacepède, 1803
Doras Lacepède, 1803:116 [type species: Silurus carinatus Linnaeus, 1766, by subsequent designation of Bleeker, 1862:5]. Gender masculine.
Mormyrostoma Miranda Ribeiro, 1911:192 [type species: Silurus carinatus Linnaeus, 1766, by original designation and monotypy]. Gender neuter.
Diagnosis.—Modern Doras is distinguished from among Doradidae by the unique combination of: 1) maxillary barbels long and fimbriate, usually reaching ventralmost opening of gill slit (Fig. 1A); 2) mesethmoid with anterior lateral margins converging towards pointed tip (Fig. 1B); 3) single anterior cranial fontanel contained largely within frontals and anteriorly by mesethmoid (Fig. 1B); 4) anterior nuchal plate wide, subpentaganol to subhexagonal, sharing broad lateral suture with epioccipital and isolating supraoccipital from middle nuchal plate (Fig. 1B); 5) nuchal foramina absent (Fig. 1B), coracoid process short, not extending much beyond posterior base of pectoral fin with tip finishing well short of that of postcleithral process; 6) dentaries typically with acicular teeth; 7) skin beneath postcleithral process perforated with conspicuous pores (Figs. 2B–H).
Sabaj Pérez et al. (2007:164) presented a diagnosis of Doras inclusive of their new fossil species, †D. dioneae, based solely on a combination of three characteristics exhibited by the postcleithral process (Fig. 2; Table 1): 1) process blade-like, subrectangular (truncated) with dorsal and ventral margins nearly parallel, posterior margin straight, weakly oblique (tilted anteriorly), and dorsal posterior corner distinct, 2) surface ornamentation separable into three longitudinal fields (dorsal, middle, ventral) with dorsal and middle fields nearly planar, and 3) middle field narrowly triangular with fine, elongate ridges and shallow grooves diverging gradually from point posterior to shoulder bulge to posterior margin of process. Their alternative diagnosis is largely corroborated by this study except the dorsal posterior corner of the postcleithral process is sometimes less distinct, broadly rounded in one species (D. phlyzakion).
Comparisons.—Fimbriate maxillary barbels (Fig. 1A) distinguish Doras from all other nominal doradid genera except Anduzedoras, Hassar, Hemidoras, Leptodoras, Nemadoras, Opsodoras and Trachydoras. The wide anterior nuchal plate sharing suture with epioccipital and absence of nuchal foramina in Doras (Fig. 1B) distinguish it from Anduzedoras, Hassar, Hemidoras, Nemadoras, Opsodoras, and Leptodoras (except L. copei and L. hasemani, from which Doras is distinguished by lacking modifications of the oral hood; see Sabaj, 2005). The short coracoid process and presence of conspicuous pores in skin beneath postcleithral process (Figs. 2B–H) distinguish Doras from three fimbriate-barbel species (Doras fimbriatus, D. punctatus, Oxydoras eigenmanni) that are incertae sedis in Doradidae (Sabaj & Ferraris, 2003). The porous skin beneath the postcleithral process in Doras is a condition similarly found only in Hassar, Nemadoras (except N. elongatus), Opsodoras ternetzi and most Trachydoras.
Doras is clearly distinguished from Trachydoras by the shape of the mesethmoid. In Doras the anterior lateral margins of the mesethmoid converge to a relatively sharp and finely notched point that finishes well beyond the first infraorbitals (lacrimals; Fig. 1B). In Trachydoras the mesethmoid is broadly rounded and expanded by lateral cornua anteriorly, and its anteriormost margin is continuous with those of the first infraorbitals.
Description.—Modern species are medium-sized doradids (to about 550 mm SL) with moderately deep and weakly compressed head and body, relatively small and subterminal mouth, large dorsal lateral eyes, and a conical snout that ranges from short and broad (D. phlyzakion, D. zuanoni) to extremely long and tapered (D. micropoeus, sometimes D. carinatus). Ventral surfaces flattened from mouth to vent. Small pores often in skin surrounding vent and in some species (D. phlyzakion, D. zuanoni) also present on abdomen and breast (Fig. 3). Infraorbital one (lacrimal) variable (Fig. 4), either short, not extended anteriorly beyond concavity in mesial margin for anterior nare (D. phlyzakion, D. zuanoni) or extended anteriorly well beyond concavity as long attenuate wing alongside autopalatine (D. carinatus, D. higuchii, D. micropoeus). Midlateral sides with 30–36 scutes beginning with infranuchal except in specimens of D. micropoeus from Maroni basin that lack anteriormost postinfranuchal scutes. Infranuchal scute (Fig. 5) tall; dorsal wing with long slender dorsal extension broadly contacting slender ventral extension of posterior nuchal plate; ventral wing shorter, lamellar, and broadly expanded anteriorly where it contacts or closely underlies distal margin of medial face of postcleithral process; medial thorn usually present and flanked by subtriangular, posteriorly pointed lamellar extensions (Figs. 5A–B) except in D. micropoeus wherein the medial thorn is absent or rudimentary and the posterior expansions are entirely lacking (Fig. 5C). All postinfranuchal scutes also usually with medial thorn and subtriangular dorsal and ventral wings (Figs. 5A–B) except in D. micropoeus wherein scutes become progressively more reduced and are sometimes lacking anteriorly. Gas (swim) bladder (Fig. 6) cordiform with broad anterior chamber contacting Müllerian rami and slightly longer, narrower posterior chambers divided internally by T-shaped septum (Fig. 6A); posteriorly with singular terminal diverticulum (D. carinatus, D. higuchii, D. micropoeus; Figs. 6A–F) or paired diverticulae separate and subterminal (D. phlyzakion; Fig. 6G) or cojoined at their bases and terminal (D. zuanoni; Fig. 6H).
Distribution and habitat.—Modern species of Doras are known from major drainages along the Atlantic coast of the Guianas from the Essequibo to the Oyapock, central Amazon drainages in northwestern Brazil (Solimões, Negro, Branco) and Colombia (Apaporis), lower Amazon drainages in northcentral Brazil (Trombetas, Jari, Xingu, Araguaia), and a southern tributary of the lower Orinoco (Paragua–Caroní) in southeastern Venezuela (Fig. 7). Doras is notable for inhabiting both lowland floodplain lakes and waterways along the Solimões-Amazon river and its tributaries (e.g., D. phlyzakion, D. zuanoni) and sizeable upland rivers on the Guiana and Brazilian Shields (D. carinatus, D. higuchii, D. micropoeus). Doras is conspicuously absent from deepwater trawls (i.e., Calhamazon collections) in both the main channels of the Amazon and lower courses of its major tributaries.
Etymology.—The origin of generic name Doras and henceforth the family Doradidae is oddly ambiguous. Lacepède’s (1803:116) descriptions of Doras and its two nominal species, D. carinatus and D. costatus, did not specify a source. An etymology for “doras” is found, however, in Lacepède’s treatment of Corydoras, a callichthyid genus that he newly proposed later in the same work. Lacepède (1803:148) explicitly noted the Greek “Corys” and “doras” to signify “casque” (helmet) and “cuirasse” (armor), respectively, in reference to the hard bony plates described separately for the head and body.
Transliterated Greek words that resemble doras and pertain specifically to armor are lacking. As a result subsequent ichthyologists have ventured to new meanings via quasi-doras transliterations of implicit Greek stems. Valenciennes (in Cuvier and Valenciennes, 1840:199 Strausbourg edition), for instance, stated without doubt that Lacepède’s Doras was derived from the Greek “dory” meaning “lance”. Miranda-Ribeiro (1911:199) later agreed. Jardine (in Schomburgk 1841:155–156) similarly speculated Doras to be from the Greek “dorat”, a small spear, and commented on the spear-shaped postcleithral process in P. costatus, a structure briefly noted by Lacepède (1803:118). Another possibility is the Greek doris, a sacrificial knife (definition from Jaeger 1950:85). Alternatively, Ringuelet et al. (1967:283) interpreted Doras as “piel, pellejo [skin, hide]” from the Greek “dora” (translated definition from Jaeger 1950:84).
Lacepède (1803:118–119) certainly appreciated the well-armed and armored nature his two species of Doras. He discussed at length the dangers that the serrated dorsal and pectoral-fin spines posed to fisherman as they removed this fish (presumably costatus, now in Platydoras) from their nets. He noted observations by Dutch naturalist Willem Pison that Brazilian fisherman believed the fin spines to be highly venomous, causing death in 24 hours unless treated with large quantities of its liver oil. Lacepède referred to the fin spines both as “dards” (meaning dart, sting or harpoon) and as “armes” (weapons) whose “dentelures” (serrations) were capable of making deep cuts in those who mishandled it.
One might consider the origin of Doras to be inspired by the catfishes’ injurous armor, technically derived from the Greek dora for animal hide, and serendipitously akin to transliterated Greek words for spear (dorat, doris, dory) in reference to the so-shaped postcleithral process and serrated fin spines.
Doras carinatus (Linnaeus, 1766)
Figs. 1A, 2B–D, 4B–C, 5A–B, 6A–B, 7, 8 & 9; Tables 1 & 2
Silurus carinatus Linnaeus, 1766:504 [type locality: Surinami (= Surinam)].
Neotype.—ANSP 187114 (alc, 155 mm), Suriname: Sipalawini: Lawa River, base camp ca. 8 km south-southwest of Anapaike/Kawemhakan (airstrip), 03°19'31”N, 054°03'48”W (SUR 07-01), M.H. Sabaj et al., 18 Apr 2007.
Non-type material.—French Guiana (Maroni Dr.): MHNG 2622.071 (3 alc, 114–142 mm), Inini River at confluence of small and large Inini rivers, Nov 1993; MHNG 2628.075 (4 alc, 41.4–68.8 mm), Tampoc River, Saut Pièrkuru, M. Jégu et al., 13 Oct 2000; MNHN 1998-1775 (1 alc, 184 mm), Tampoc River, St. Laurent du Maroni State, P.-Y. Le Bail & P. Keith, Nov 1998; MNHN 2000-4468 (3 alc), Maroni River, Maripasoula, 15 Jun 1999; MNHN 2000-5863 (1 of 2), Tampoc River, Saut Pièrkuru (station niv1mar4), 02°49'N, 053°32'W, M. Jégu et al., 2000; French Guiana (Sinnamary Dr.): MNHN 1998-1820 (3 alc 181-218 mm) Sinnamary River, 10 km in front of Sinnamary, Cayenne State, P.-Y. Le Bail, 25 May 1981; French Guiana (Oyapock Dr.): MNHN 1998-1693 (1, 196 mm), Oyapock basin (station d), Cayenne State, P.-Y. Le Bail & P. Keith, 19 Oct 1986; MNHN 1981-0247 (2 alc), Crique Pakoti, Cayenne State, Mr. Grenand, 11 Sep 1976; MNHN 1981-0257 (1), Oyapock River, Trois sauts, D'Aubenton et al., 3 Oct 1976; MHNG 2681.006 (1 alc, 154 mm), Oyapock River (main channel), downstream of Moulou Koulou creek, Camopi, Alicoto, 03°06'750”N, 052°20'463”W, R. Covain et al., 2006; MHNG 2681.071 (1 alc, 168 mm), Oyapock River (main channel), upstream of Fifine creek at Wacarayou rapids, 03°13'672”N, 052°17'577”W, R. Covain et al., 2006. Guyana (Essequibo Dr.): AMNH 7069 (3 alc, 72-136.5 mm), Essequibo River, Crabb Falls, C.H. Eigenmann, S.E. Shideler et al., 4–7 Nov 1908; AMNH 17632 (1 alc, 188 mm), upper Essequibo River, on or above cataract, T. Holden Expedition, 28 Dec 1937; AMNH 214981 (12 alc, 49.5-99.2 mm), Essequibo River, A.S. Pinkus, 1935; ANSP 175870 (1 alc, 65.9 mm), Essequibo River, sandbars in vicinity of Maipuri campsite, 04°34'17”N, 058°35'17”W (WGS97-28), W.G. Saul et al., 31 Jan 1997; ANSP 175871 (1 alc, 55.3 mm), Essequibo River at Essequibo campsite, 04¯45'41^N, 058¯45'53^W (WGS97-19), D. Torres et al., 26 Jan 1997; ANSP 175872 (4 alc, 42.9–52.3 mm), Essequibo River, sandbars in vicinity of Maipuri campsite, 04°34'17”N, 058°35'17”W (WGS97-31), W.G. Saul et al., 2 Feb 1997; ANSP 175873 (2 alc, 46–47.9 mm), Isolated stagnant pool/pond some 40 minutes from main Essequibo River channel, 04°32'43”N, 058°35'02”W (WGS97-27), W.G. Saul et al., 31 Jan 1997; ANSP 177272 (1 alc, 120.2 mm), Burro Burro River, creek tributary downstream from Burro Burro camp-between Lunch Spot and Water Dog Camp, 04°41’N, 058°51’W (GGW97-10), G. Watkins et al., 20 Nov 1997; ANSP 177273 (6 alc, 107.3–153.6 mm), Essequibo River, extensive sandbar 2.0 km upstream from Paddle Rock campsite, 04°42'20”N, 058°42'26”W (GGW97-23), C. Watson et al., 25 Nov 1997; ANSP 177274 (2 alc, 89.6–95.7 mm), Essequibo River, extensive sandbar 500m downstream from Paddle Rock campsite, 04°44'00”N, 058°43'00”W (GGW97-17), C. Watson et al., 23 Nov 1997; ANSP 177275 (1 alc, 188.0 mm), Siparuni River, Blackwater camp and blackwater creek, 04°44'21”N, 058°57'54”W (GGW97-27), G. Watkins et al., 4 Dec 1997; ANSP 177276 (17 alc, 82.1–196 mm), Essequibo River, Yurrie Creek approx. 2.0 km upstream from Paddle Rock campsite, 04°42'03”N, 058°42'44”W (GGW97-24A), C. Watson et al., 26 Nov 1997; ANSP 178704 (1 alc, 97.4 mm), Essequibo River, 180 yd. upstream from Essequibo campsite (Maipuri), 04°45'43”N, 058°45'52”W (WGS97-23), D. Allicock, 27 Jan 1997; ANSP 179619 (3 alc, 33.6–47.4 mm), Essequibo River (east bank) at Kurukupari, 04°39'41”N, 058°40'31”W (GUY 02-01), M.H. Sabaj et al., 24 Oct 2002; ANSP 180986 (1 sk, 170 mm), Essequibo River, Yukanopito Falls, 44.5 km SW of mouth of Kuyuwini River, 01°54'53”N, 058°31'14”W (GUY 03-19), M.H. Sabaj et al, 9 Nov 2003; ANSP 182334 (2 alc, 34.2–47.3 mm), Essequibo River, Kassi-Attae Rapids, 5.5 km SE of mouth of Kuyuwini River, 02°13'36”N, 058°17'38”W (GUY 03-15), M.H. Sabaj et al, 8 Nov 2003; ANSP 185199 (1 alc, 108.5 mm), Kuyuwini River, 60.6 km ENE of Kuyuwini Landing, 179 km SE of Lethem, 02°11'35”N, 058°42'15”W (GUY 03-11), M.H. Sabaj et al, 6 Nov 2003; ANSP 185200 (1 alc, 36.2 mm), Kuyuwini River, main channel and backwater 19.5 km W of confluence with Essequibo River, 02°14'28”N, 058°30'03”W (GUY 03-22), M.H. Sabaj et al, 11 Nov 2003; AUM 27845 (1 alc, 125.5 mm), Essequibo River, Rockstone, 05°59'07.5"N, 058°33'02.9"W (Guy 98-14), M.H. Sabaj et al., 19–20 Oct 1998; AUM 28013 (1 alc, 47.4 mm), Essequibo River, large sandbar & small cataract, 31.9 mi SSW Rockstone, bearing 204°, 05°31'39.5"N, 058°37'43.6"W (Guy 98-17), J.W. Armbruster et al., 21 Oct 1998; FMNH 53189 (1 alc, 74.7 mm), Bartica, C.H. Eigenmann et al., 1908; FMNH 53190 (10 alc, 57.5–194 mm), Rockstone, C.H. Eigenmann et al., 1908; FMNH 53191 (1 alc, 36.5 mm), 53734 (1 alc, 207.5 mm), Georgetown Market, C.H. Eigenmann, et al., 1908; FMNH 53711 (72 alc), Potaro River at Tumatumari, C.H. Eigenmann, et al., 1908; INHS 49235 (1 alc), Essequibo River, 0.72 mi SW Rockstone at sandbar, bearing 221°, 05°58'34.0"N, 58°33'19.3"W (Guy 98-11), M.H. Sabaj et al., 19 Oct 1998; INHS 49321 (2 alc, 141–152 mm), same data as AUM 27845; INHS 49359 (2 alc, 40.3–131.7 mm), same data as AUM 28013; INHS 49445 (43 alc, 31–43.2 mm), Potaro River, beach on N bank, downstream of Tumatumari Cataract, 05°21'48.4"N, 059°00'04.4"W (Guy 98-20), M.H. Sabaj et al., 22 Oct 1998; MZUSP 88605 (4 alc, 77.2–157.7 mm), same data as ANSP 177276; ROM 62641 (13 alc, 31.5–50.6 mm), Essequibo River, inlet and beach downstream from Kurupukari, 04°42'57"N, 058°42'40"W (H90-43), E. Holm et al., 10 Oct 1990; USNM 66201 (4 alc, 47.9–83.8 mm), same data as FMNH 53711; USNM 66202 (6 alc, 36.8–134.9 mm), Essequibo River at Crab Falls, C.H. Eigenmann, et al., 1908.