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W. M. N. -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Vertebrate Section, Ichthyology, Royal Museum for Central Africa (RMCA), Tervuren, Belgium - - - -Author - -Bragança Tobias Musschoot, Pedro H. N. - - - -Author - -Chakona, Albert - -text - - -Zoological Journal of the Linnean Society - - -2024 - -202 - - -1 -33 - - - -journal article -10.1093/zoolinnean/zlae121 -0024-4082 -14544245 -3C2308D-7334-412D-871F-DE1F17C38D0E - - - - - - - -Parauchenoglanis ngamensis -( -Boulenger 1911 -) - - - - - - - - -( -Fig. 5 -; -Table 3 -) - - - - - - -Auchenoglanis ngamensis -Boulenger 1911 - -. - - - -Parauchenoglanis ngamensis - -( -Mo 1991 -, - -Teugels -et al. -1991 - -, -Skelton 2001 -, -2019 -, - -Geerinckx -et al. -2003 - -, -2004 -, -Ferraris 2007 -, -Retzer 2008 -, -Marshall 2011 -, - -Bruton -et al. -2018 - -). - - - -Parauchenoglanis ngamensis - -‘okavango–zambezi’ ( - -Sithole -et al. -2023 - -). - - - - -Common English name: -Zambezi -grunter ( -Skelton 2001 -). - - -Common French name: -Mâchoiron de la -Zambezi -. - - - - -Holotope: - - -BMNH 1910.5.31.35 - -, -184.5 mm -SL, -Botswana -, -Okavango Basin -, near -Lake Ngami -(~ - -20°23 -ʹ -42.0″S - -, - -22°55 -ʹ -32.9″E - -); collector -R.B. Woosnam -1911 - -. - - -Non-tope material (N = 62): - -Angola - -: -RMCA -_ -Vert -_P.161712−161713, two, -139.6–204.2 mm -SL, -upper Zambezi -sub-basin, -Lake Calundo -( - -11°48 -ʹ -S - -, - -20°52 -ʹ -E - -); collector unspecified; 1964. -RMCA -_ -Vert -_P.161715, one, 169.0 mm SL, -upper Zambezi -sub-basin, -Lake Calundo -( - -11°48 -ʹ -S - -, - -20°52 -ʹ -E - -); collector unspecified; 1964. -SAIAB 190829 -, one, -41.1 mm -SL, -upper Zambezi -sub-basin, -Cuando Cubango -, -Jamba -camp site 2 ( - -17°32 -ʹ -4.4″ -S - -, - -23°11 -ʹ -20.7″ -E - -); collectors -Nkosinathi Mazungula -, -Manuel Domingos -and -Francisco de Almeida -; - -27 April 2013 - -. -SAIAB 207888 -, two, 24.0– -89.5 mm -SL, -Okavango Basin -, -Rio Comba -(affluent of -Lungwebungu -) -Sassicola Village -( - -13°08 -ʹ -12.8″ -S - -, - -19°01 -ʹ -48.3″ -E - -); collectors -Paul Skelton -, -Ben C.W. van der Waal -and -Frowin Becker -; - -22 April 2018 - -. -SAIAB 203307 -, one, -104.2 mm -SL, -Okavango Basin -, -Cuito -, -Samahamba Village -( - -14°51 -ʹ -17″ -S - -, - -19°17 -ʹ -11″ -E - -); DNA sample PHS16-335: SB8972; collector -Adjany Costa -; - -1 March 2016 - -. -SAIAB 101057 -, two, -121.9–134.4 mm -SL, -Okavango Basin -, -Cuito -, margins of main channel, protected cattle drinking area and swamp ( - -17°2 -ʹ -55.4″ -S - -, - -19°32 -ʹ -4.6″ -E - -); collectors -Roger Bills -, -Nkosinathi Mazungula -and -Manuel Domingos -; - -17 July 2013 - -. -SAIAB 203213 -, four, 112.3–147.0 mm SL, -Okavango Basin -, -Cunde Waterfall -( - -13°46 -ʹ -25.0″ -S - -, - -18°45 -ʹ -22.8″ -E - -); collector -Adjany Costa -; - -2 March 2016 - -. -SAIAB 186847 -, three, -72.8–132.9 mm -SL, -Okavango Basin -, -Cuito -, -Cunavale -bridge ( - -15°10 -ʹ -16.4″ -S - -, - -19°11 -ʹ -39.3″ -E - -); DNA sample RB12-A155: -ANGFW201-12 -; collectors -Roger Bills -, -Paul Skelton -and -Francisco de Almeida -; - -18 May 2012 - -. -SAIAB 203177 -, one, -24.4 mm -SL, -Okavango Basin -, -Cuanavale Source Lake -( - -13°05 -ʹ -25.1″ -S - -, - -18°53 -ʹ -37.4″ -E - -); collector -Adjany Costa -; - -28 February 2016 - -. -SAIAB 186832 -, four, -16.9– 53.1 mm -SL, -Okavango Basin -, -Luassingua River -, -55 km -east of -Menongue -, at road -Menongue-Cuito Cuanavale -( - -14°35 -ʹ -24.2″ -S - -, - -18°10 -ʹ -14.4″ -E - -); DNA sample RB12-A474: -ANGFW168-12 -; collector -Roger Bills -; - -17 May 2012 - -. -SAIAB 202475 -, one, -84.4 mm -SL; -Okavango Basin -, -Hippo -camp ( - -13°08 -ʹ -29.0″ -S - -, - -18°29 -ʹ -08.0″ -E - -); collector -Adjany Costa -; - -11 June 2015 - -. -SAIAB 204028 -, four, -46.2–111.6 mm -SL, -Okavango Basin -, below the outlet of the -Cuanavale Source Lake -( - -13°05 -ʹ -37.1″ -S - -, - -18°53 -ʹ -45.5″ -E - -); collectors -Ben C.W. van der Waal -and -Nkosinathi Mazungula -; - -23 October 2016 - -. -SAIAB 203160 -, two, -36.6–45.2 mm -SL, -Okavango Basin -, -Kalva -and -Cuito Confluence -( - -12°44 -ʹ -55.6″ -S - -, - -18°21 -ʹ -15.6″ -E - -); collector -Adjany Costa -; - -25 February 2016 - -. -SAIAB 203267 -, two, -104.6–113.7 mm -SL, -Okavango Basin -, -Cuito -, -Alexandra -camp ( - -14°24 -ʹ -40.1″ -S - -, - -18°56 -ʹ -54.5″ -E - -); collector -Adjany Costa -; - -26 June 2015 - -. -SAIAB 186951 -, two, -42.9 mm -SL, -Okavango Basin -, -Cuchi town -, road bridge ( - -14°38 -ʹ -58.9″ -S - -, - -16°54 -ʹ -23.7″ -E - -); collectors -Roger Bills -and -Paul Skelton -; - -21 May 2012 - -. -SAIAB 190653 -, three, -174.1–198.5 mm -SL -, -upper Zambezi -sub-basin, -Cuando Cubango -, -Base-Rio -campsite ( - -17°34 -ʹ -23.1″ -S - -, - -23°15 -ʹ -36.9″ -E - -); collectors -Nkosinathi Mazungula -, -Manuel Domingos -and -Francisco de Almeida -; - -30 April 2013 - -. -SAIAB 204289 -, three, -36.3−125.9 mm -SL, -upper Zambezi -sub-basin, -Crossing -on maintrack -SW Cambuta -( - -13°26 -ʹ -48.4″ -S - -, - -19°57 -ʹ -50.5″ -E - -); DNA sample -ANG16-306 -: SB 7581; collector -Roger Bills -; - -20 November 2016 - -. -SAIAB 204017 -, two, -62.2–74.1 mm -SL, -upper Zambezi -sub-basin, -Lungwebungu River -near bridge ( - -12°34 -ʹ -54.5″ -S - -, - -18°40 -ʹ -09.5″ -E - -); collectors -Ben C. W. van der Waal -and -Nkosinathi Mazungula -; - -20 October 2016 - -. -SAIAB 204255 -, one, -112.7mm -SL, -upper Zambezi -sub-basin, -Cuembe -at road bridge ( - -13°31 -ʹ -35.1″ -S - -, - -19°16 -ʹ -47.6″ -E - -); DNA sample -ANG16-338 -: SB7582; collector -Roger Bills -; - -19 November 2016 - -. - -Botswana - -: -SAIAB 18928 -, one, -44.8 mm -SL, -Okavango -Basin -, -Lake Ngami -, -East -shore ( - -20°30 -ʹ -0″S - -, 22°40 -ʹ -.1″E); collector -Mike N. Bruton -; - -7 June 1980 - -. -SAIAB 52432 -, one, -149.1 mm -SL, -Okavango -Basin -, -Shakowe Camp -( - -18°25 -ʹ -59.9″ -S - -, - -21°54 -ʹ -00.0″ -E - -); collectors -Roger Bills -and -P. Yose -; - -26 May 1996 - -. - -Namibia - -: -SAIAB 187013 -, one, -40.3 mm -SL, -Okavango -Basin -, -Popa -, ( - -18°07 -ʹ -18.2″ -S - -, - -21°35 -ʹ -00.2″ -E - -); DNA sample RB12-540: -ANGFW222-12 -; collectors -Roger Bills -and -Paul Skelton -; - -24 May 2012 - -. -SAIAB 203080 -, one, -102 mm -SL, -Okavango -Basin -, -River -dance camp ( - -19°54 -ʹ -52.4″ -S - -, - -24°22 -ʹ -50.2″ -E - -); collector -Adjany Costa -; - -5 August 2015 - -. -SAIAB 202381 -, three, -31.6–145.7 mm -SL, -Okavango -Basin -, -Carioo -unloading camp ( - -12°52 -ʹ -31.0″ -S - -, - -18°18 -ʹ -6.0″ -E - -); collector -Paul Skelton -; - -3 June 2015 - -. -SAIAB 44835 -, one, -104.6 mm -SL, upper -Zambezi -sub-basin ( - -17°28 -ʹ -00.0″ -S - -, - -24°17 -ʹ -00.0″ -E - -); collector -C. Hay -; - -1 May 1994 - -. -SAIAB 85562 -, one, -83.9 mm -SL, upper -Zambezi -sub-basin ( - -17°28.5 -ʹ -S - -, - -24°14.9 -ʹ -E - -); DNA sample RB08 -ZaNam -281: SB7585; collector -Roger Bills -; - -29 September 2008 - -. - -Zambia - -: -RMCA -_ -Vert -_P.142057, one, -137.3 mm -SL, -upper Zambezi -sub-basin, -Chavuma -, -Balovale -( - -13°5 -ʹ -S - -, - -22°40 -ʹ -E - -); collector -G. Bell-Cross -; 1963. -RMCA -_ -Vert -_P.142072-142073, one, -101.4 mm -SL, -upper Zambezi -sub-basin, -Monga -( - -15°17 -ʹ -S - -, - -22°50 -ʹ -E - -); collector -G. Bell-Cross -; 1964. -SAIAB 98049 -, three, -57.2–131.8 mm -SL, -upper Zambezi -sub-basin, -Kalumbila -stream on -K Hill -road ( - -12°14 -ʹ -34.4″ -S - -, - -25°22 -ʹ -43.2″ -E - -); DNA sample JWH10-A162: SB7586; collectors -Roger Bills -and -A. Bok -; - -8 November 2010 - -. -SAIAB 71358 -, two, -88.7−105.8 mm -SL, -upper Zambezi -sub-basin ( - -15°11 -ʹ -43″ -S - -, - -22°55 -ʹ -20″ -E - -); collectors -Denis Tweddle -and -Ben C.W. van der Waal -; - -10 November 2002 - -. -SAIAB 97996 -, one, -22.8 mm -SL, -upper Zambezi -sub-basin, -Mupulumba Village -at footbridge ( - -12°02 -ʹ -28.8″ -S - -, - -25°25 -ʹ -33.9″ -E - -); collectors -Roger Bills -and -A. Bok -; - -4 November 2010 - -. -SAIAB 71869 -, one, -97.9mm -SL, -upper Zambezi -sub-basin ( - -15°22 -ʹ -29″ -S - -, - -23°12 -ʹ -2″ -E - -); collectors -Denis Tweddle -and -Ben C.W.van der Waal -; - -27April 2003 - -. -SAIAB 85560 -, one, -47.2 mm -SL, -upper Zambezi River -, -Katima Mulilo -in front of -Ben’s House -( - -17°29 -ʹ -11″ -S - -, - -24°15 -ʹ -25″ -E - -); DNA sample RB08 -ZaNam -187: SB8974; collector -Roger Bills -; - -29 September 2008 - -. -SAIAB 98109 -, one, -114.2 mm -SL, -upper Zambezi -sub-basin, -Enterprise -mine camp ( - -12°11 -ʹ -29.3″ -S - -, - -25°12 -ʹ -31.8″ -E - -); DNA sample JWH10-A076: SB7588; collectors -Roger Bills -and -A. Bok -; - -6 November 2010 - -. -SAIAB 98168 -, two, -52.5–89.6 mm -SL, -upper Zambezi -sub-basin, -Chisolo river -above -Enterprise -camp on drilling road (west bank) ( - -12°10 -ʹ -31.8″ -S - -, - -25°12 -ʹ -54.8″ -E - -); DNA sample JWH10-A156: SB7587; collectors -Roger Bills -and -A. Bok -; - -6 November 2010 - -. -SAIAB 73614 -, one, -137.2 mm -SL, supper -Zambezi -sub-basin ( - -13°25 -ʹ -21″ -S - -, - -24°19 -ʹ -54″ -E - -); collectors -Denis Tweddle -and -Ben C. W. van der Waal -; - -6 October 2003 - -. - - - - -Table 2. -Comparison of qualitative characters between the - -Parauchenoglanis ngamensis - -candidate species identified based on the -COI -sequence data and the two candidate species identified by the morphological data only. - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
-Character - -Clade 1 - -Clade 2 - -Clade 3 - -Morphological data only -
- -P. ngamensis -s.s. - - - -P. ngamensis - - - -P. ngamensis - - - -P. ngamensis - - - - -P. ngamensis - -sp. - - - - -P. ngamensis - -sp. - - - - -P. ngamensis - -sp. - - - - -P. ngamensis - -sp. - - - - -P. ngamensis - -sp. - -
-sp. ‘kwanza 1’ - - -sp. ‘kwanza 2’ (= -P. - - -sp. ‘kasai 1’ - -‘kasai 2’ - - -‘kasai 3’ (= -P. - - -‘kasai 4’ - -‘luenda’ - - -‘chiumbe’ (= -P. - -
- -(= - -P. patersoni - -) - - - -dolichorhinus -) - - - -(= - -P. lueleensis - -) - - - -(= - -P. poikilos - -) - - - -ernstswartzi -) - - - -(= -P. - - - -(= - -P. luendaensis - -) - - - -chiumbeensis -) - -
- -megalasma -) - -
Caudal finRoundRoundRoundTruncatedTruncatedTruncatedTruncatedTruncatedTruncated
Colouration
BodyBars (<100 mmBars (<65 mmBars (<95 mm SL),SpottedSpottedBlotchesBlotchesSpottedSpotted
SL), spotsSL), spotsspots (>100 mm
(≥100 mm SL)(>100 mm SL)SL)
Back -Numerous spotsAbsentAbsentOne or twoNumerous spotsVermiculatedBlotchesAbsentAbsent
ground*spotspattern
HeadNumerous spotsAbsent (<10 ifAbsent (<10 if pre-Absent (<10 ifNumerous spotsBlotchesBlotchesAbsentFaint spots
present)sent)present)
Head profileModerately de-Moderately de-DepressedModerately de-Moderately de-Moderately de-Moderately de-Moderately de- -Moderately -
depressedpressedpressedpressedpressedpressedpressedpressed
Snout profileBluntly triangularBluntly triangularBluntly triangularBluntly tri-Bluntly tri-RoundPartly roundBluntly triangularBluntly triangular
angularangular
Eyes profileDorsolateralDorsolateralDorsolateralDorsolateralDorsolateralDorsalDorsolateralDorsolateralDorsolateral
- - - -* -Background = additional black spots between vertical rows of spots or blotches. - - - - - -Diagnosis: -Parauchenoglanis ngamensis - -is distinguished from - -Parauchenoglanis ahli -(Holly 1930) - -, - -Parauchenoglanis altipinnis -( -Boulenger 1911 -) - -, - -P. balaoi -(Sauvage 1879) - -, - -Parauchenoglanis buetikoferi -(Popta 1913) - -, - -Parauchenoglanis longiceps -(Boulenger 1913) - -, - -P. monkei -(Keilhack 1910) - -, - -Parauchenoglanis pantherinus -(Pellegrin 1929) - -, and - -P. punctatus - -by a broad humeral process (vs. pointed humeral process). It is further distinguished from - -P. altipinnis - -, - -P. balaoi - -, - -P. pantherinus - -, and - -P. punctatus - -by coarse skin (vs. smooth skin). It is also further distinguished from - -P. buetikoferi - -and - -P. monkei - -by anterior margin of the pectoral-fin spine mostly smooth (vs. anterior margin of pectoral-fin spine serrated from base to tip). It is further distinguished from - -P.ahli - -and - -P. punctatus - -by shorter barbels, with external mandibular barbel not reaching the distal tip of the pectoral fin (vs. barbels long, with external mandibular barbel reaching distal tip of pectoral fin or beyond). - -Parauchenoglanis ngamensis - -is distinguished from - -P. zebratus - -by broad humeral process clearly visible through the skin and presence of five to seven bars on small specimens (≤ -100 mm -SL) and vertical rows of spots on larger-sized specimens [vs. broad humeral process embedded under the skin and four to five vertical rows of spots on small specimens (≤ -100 mm -SL) and bars on larger-sized specimens]. - -Parauchenoglanis ngamensis - -is distinguished from - -P. stiassnoae - -by the presence of vertical rows of black spots interspaced by scattered smaller black spots (vs. irregularly scattered large blotches and absence of smaller spots on the flank). They are further distinguished by shorter barbels, external mandibular barbel not reaching the distal tip of the pectoral-fin spine in - -P. ngamensis - -(vs. longer barbels, external mandibular extending beyond tip of the pectoral-fin spine in - -P. stiassnoae - -). - - - -Figure 5. -Photographs of the holotype of - -Parauchenoglanis ngamensis -, -BMNH - -1910.5.31.35 -, 184.5 mm SL, Okavango River. A, lateral view. B, dorsal view. C, ventral view. Scale bar: 1 cm. - - - - -Parauchenoglanis ngamensis - -is distinguished from the other members of the - -P. ngamensis - -group, i.e. - -P. patersoni - -, - -P. dolichorhinus - -, - -P. luendaensis - -, and - -P -. -chiumbeensis - -, by the presence of scattered black spots in between the vertical rows of black spots (vs. absent). It is further distinguished from these species by heavily spotted head and fins (vs. unspotted head and fins in - -P -. -luendaensis - -; few black spots or unspotted head and fins in - -P -. -patersoni - -and - -P. dolichorhinus - -, and head and fins with faint black spots in - -P. chiumbeensis - -). - -Parauchenoglanis ngamensis - -is further distinguished from - -P -. -chiumbeensis - -and - -P. luendaensis - -by round caudal fin ( -Fig. 5A -; vs. truncated, -Figs 14A -, -15A -, respectively). - -Parauchenoglanis ngamensis - -is distinguished from - -P -. -ernstswartzi - -and - -P -. -megalasma - -by vertical rows of black spots ( -Fig. 5A -; vs. vertical rows of black blotches, -Figs 12A -, -13A -, respectively) and bluntly triangular snout ( -Fig. 5B -; vs. round in - -P. ernstswartzi - -and partly round in - -P. megalasma - -, -Figs 12B -, -13B -, respectively). It is further distinguished from - -P -. -ernstswartzi - -by eyes situated dorsolateral ( -Fig. 5A -; vs. eyes situated dorsally, i.e. high on the head towards its upper edge, -Fig. 12A -) and absence of black spots at the base of pectoral fin ( -Fig. 5C -; vs. present, -Fig. 12C -). - -Parauchenoglanis ngamensis - -is distinguished from - -P. lueleensis - -by presence of numerous black spots in between vertical rows ( -Fig. 5A -; vs. one or two spots present in between vertical rows, -Fig. 12A -). Finally, - -P. ngamensis - -is distinguished from - -P. poikilos - -by a rounded caudal fin ( -Fig. 5A -; vs. truncated, -Fig. 11A -) and the absence of crescent-shaped marking on the caudal-fin base ( -Fig. 5A -; vs. present, -Fig. 11B -). - - - - -Table 3. -Morphometrics and meristics of the - -Parauchenoglanis -species - -(re)described in this study.The given range for a species also includes holotype and paratype(s). - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- - -P. ngamensis -s.s. - -( -N -= 63) - - - - -P. -patersoni - -( -N -= 14) - - - - -P. dolichorhinus - -( -N -= 4) - -
-Variables - -Holotype - -Range - -Mean - -Holotype - -Range - -Mean - -Holotype - -Range - -Mean -
Standard length (mm)184.522.8–204.298.8192.942.9–223.0111.5139.852.4–115.186.5
Body measurements (% standard length)
Preanal length60.955.9–74.164.061.156.1–62.260.658.858.6–59.959.4
Prepelvic length50.248.0–54.950.952.148.1–53.551.148.949.6–50.950.2
Predorsal length38.836.0–42.739.738.336.8–42.139.637.137.2–39.838.1
Prepectoral length27.425.6–31.228.426.326.7–32.528.728.126.8–28.527.6
Adipose fin length33.528.7–41.033.534.131.6–38.834.832.732.8–34.733.9
Interdorsal-adipose distance4.53.0–12.97.76.93.5–10.06.410.56.3–9.98.4
Dorsal fin base length15.914.2–18.416.315.514.7–18.716.215.515.7–17.616.5
Dorsal fin length15.213.5–25.119.916.215.3–21.018.416.215.5–21.117.8
Dorsal fin spine length14.012.4–20.316.316.614.5–19.516.311.311.4–13.712.7
Pectoral fin length17.115.9–24.020.018.414.0–22.419.118.516.1–21.218.8
Pectoral fin spine length16.712.7–21.117.016.512.9–19.016.916.612.6–15.814.6
Pelvic fin length13.111.6–17.515.115.012.2–14.913.414.511.6–14.813.3
Anal fin length12.110.2–25.116.213.310.6–14.612.312.613.2–14.213.7
Interpectoral distance14.715.9–25.019.718.417.0–23.820.218.217.4–17.817.6
Interpelvic distance6.85.2–12.08.610.98.6–11.39.911.09.8–11.210.3
Maximum body depth17.414.6–23.518.416.315.5–21.417.613.211.6–14.012.9
Pelvic body depth16.59.3–21.014.613.111.3–16.613.09.28.7–9.39.1
Minimum caudal peduncle depth11.08.4–13.611.211.79.9–12.111.19.710.0–11.810.9
Maximum caudal peduncle depth14.512.3–17.915.517.011.3–16.613.715.414.0–14.414.3
Adipose fin-caudal fin interdistance6.84.8–8.36.65.44.1–6.75.95.85.3–6.45.9
Anal fin-caudal fin interdistance14.712.6–18.515.214.112.4–17.115.217.614.0–17.716.0
Adipose fin depth4.64.6–8.26.47.75.8–8.57.34.95.0–6.25.6
Head length34.330.6–35.533.032.630.2–34.933.133.632.5–32.732.6
Head measurements (% head length)
Postorbital head length42.639.1–47.942.943.039.9–46.143.541.541.4–43.342.5
Preorbital head length45.434.3–50.945.847.238.2–50.844.048.144.8–45.245.0
Head width49.560.9–85.970.166.666.8–78.571.365.863.1–72.567.5
Orbital head width54.352.2–69.160.057.057.5–64.260.449.449.4–57.654.0
Head depth47.640.6–60.852.352.548.5–54.951.734.638.9–40.039.6
Orbital head depth33.121.8–48.136.740.827.4–45.936.925.827.6–28.628.2
Snout depth18.110.9–28.517.620.413.2–20.917.013.512.1–12.512.2
Maxillary barbel length42.730.3–79.048.137.339.4–62.050.648.360.4–72.166.1
- - - -Table 3. -Continued - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- - -P. ngamensis -s.s. - -( -N -= 63) - - - - -P. -patersoni - -( -N -= 14) - - - - -P. dolichorhinus - -( -N -= 4) - -
-Variables - -Holotype - -Range - -Mean - -Holotype - -Range - -Mean - -Holotype - -Range - -Mean -
External mandibular barbel length81.248.6–110.185.386.076.7–118.293.086.399.5–118.1106.0
Internal mandibular barbel length44.431.0–55.843.537.836.1–58.345.235.340.7–52.745.8
Mandibular barbels interdistance13.59.3–21.414.014.010.3–18.714.612.012.4–13.413.0
Interorbital distance22.019.3–28.022.620.019.7–25.822.817.516.7–22.819.7
Anterior nostrils interdistance9.47.6–20.213.09.513.7–20.316.810.57.2–17.613.4
Posterior nostrils interdistance7.58.5–12.210.08.97.9–11.19.48.88.0–9.28.5
Supraoccipital process-nuchal plate interdistance1.3.9–7.22.71.31.4–6.73.66.74.0–6.85.9
Supraoccipital process-dorsal fin interdistance18.113.9–23.217.918.210.2–21.215.114.09.2–15.013.0
Prehyoid length15.47.9–21.715.720.410.3–25.816.020.815.5–21.018.3
Mouth width41.727.6–66.646.645.740.2–61.246.947.644.0–54.448.9
Premaxillary toothplate width9.58.7–19.412.211.49.1–13.211.312.08.8–15.811.5
Orbital diameter13.111.4–24.516.515.514.2–19.716.915.716.4–19.517.5
MeristicsMedianMedianMedian
No.of dorsal-fin rays77–8776–77777
No.of anal-fin rays108–11101110–111089–109
No.of pectoral-fin rays87–88777777
No.of pelvic-fin rays666666666
No.of caudal-fin rays1815–18171815–17171415–1615
No.of serrae anterior margin pectoral-fin spine20–7352–5342–43
No.of serrae posterior margin pectoral-fin spine184–1912177–169158–109
Vertebrae -( -N -= 4) - -( -N -= 4) - -( -N -= 4) -
No.of pre-anal vertebrae1515–17151616161515–1715
No.of caudal vertebrae2019–21192019–20192019–2120
Total Vertebrae3534–36363635–3635.53535–3635.5
- - - -Table 3. -Continued - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- -P. lueleensis - -( -N -= 14) - - -P. poikilos - -( -N -= 2) - - -P. ernstswartzi - -( -N -= 4) - - -P -. -megalasma - -( -N -= 2) -
-Variables - -Holotype - -Range - -Mean - -Holotype - -Paratype - -Holotype - -Range - -Mean - -Holotype - -Paratype -
Standard length (mm)95.432.1–158.565.657.236.396.692.0–105.697.572.566.9
Body measurements (%standard length)
Preanal length62.557.0–61.359.859.660.161.161.5–65.263.559.459.5
Prepelvic length51.250.1–53.851.450.051.153.153.8–57.155.551.154.0
Predorsal length39.839.4–42.040.739.238.440.139.1–40.540.040.741.9
Prepectoral length30.028.5–30.929.831.430.629.030.3–31.831.028.630.4
Adipose fin length36.732.5–39.736.936.036.635.333.2–35.734.436.935.6
Interdorsal-adipose distance3.74.1–6.85.25.55.83.93.8–5.04.44.83.9
Dorsal fin base length14.915.5–17.816.615.816.316.616.4–17.717.116.918.2
Dorsal fin length17.514.5–23.419.319.417.619.116.4–19.918.622.921.4
Dorsal fin spine length16.712.3–16.615.119.215.614.615.8–21.218.515.016.8
Pectoral fin length20.819.2–22.521.521.820.522.515.7–22.019.121.921.6
Pectoral fin spine length17.614.6–19.817.816.914.820.218.5–18.918.716.819.7
Pelvic fin length16.011.8–17.315.416.613.415.915.0–16.115.615.315.6
Anal fin length16.710.8–15.913.916.312.014.514.6–14.814.716.217.0
Interpectoral distance18.918.8–22.520.518.421.019.919.8–20.620.218.821.9
Interpelvic distance10.07.1–11.29.18.97.69.58.3–10.39.69.39.8
Maximum body depth15.715.0–17.416.116.216.018.718.2–19.918.920.220.9
Pelvic body depth12.18.8–14.310.811.911.414.314.2–16.915.414.312.8
Minimum caudal peduncle depth12.210.6–13.011.611.210.412.313.4–14.613.913.113.4
Maximum caudal peduncle depth14.711.1–16.013.913.913.515.114.9–15.215.115.514.9
Adipose fin-caudal fin interdistance5.85.7–6.86.26.25.75.94.6–6.25.25.74.2
Anal fin-caudal fin interdistance16.714.5–17.616.417.013.117.912.7–15.113.815.015.3
Adipose fin depth7.66.1–8.17.06.46.17.37.3–7.57.57.37.8
Head length36.031.8–35.733.935.635.033.234.9–35.335.134.835.5
Head measurements (% head length)
Postorbital head length42.238.8–45.441.540.642.441.140.7–42.041.440.638.6
Preorbital head length44.437.6–47.242.442.041.247.141.9–45.644.245.746.6
Head width64.864.9–78.371.565.368.870.468.0–70.568.966.272.6
Orbital head width54.554.3–64.259.650.356.362.561.4–61.761.558.259.6
Head depth44.941.2–50.345.943.541.752.549.5–54.052.152.252.7
Orbital head depth35.921.4–36.328.727.226.740.234.2–42.038.938.538.3
Snout depth15.912.3–19.515.013.811.018.616.5–19.318.118.317.0
Maxillary barbel length59.836.1–66.756.368.360.354.553.4–60.956.554.369.7
External mandibular barbel length122.860.5–123.5102.1109.5102.289.485.2–92.989.789.699.2
- - - -Table 3. -Continued - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- -P. lueleensis - -( -N -= 14) - - -P. poikilos - -( -N -= 2) - - -P. ernstswartzi - -( -N -= 4) - - -P -. -megalasma - -( -N -= 2) -
-Variables - -Holotype - -Range - -Mean - -Holotype - -Paratype - -Holotype - -Range - -Mean - -Holotype - -Paratype -
Internal mandibular barbel length50.935.8–52.644.245.653.060.252.8–57.455.553.860.3
Mandibular barbels interdistance14.115.5–21.417.315.618.814.413.0–15.914.814.812.4
Interorbital distance17.318.0–26.221.119.224.621.418.3–20.919.321.222.6
Anterior nostrils interdistance13.315.7–21.718.018.117.819.115.1–18.216.720.022.0
Posterior nostrils interdistance8.17.9–10.99.110.111.09.910.5–11.411.110.510.9
Supraoccipital process-nuchal plate interdistance2.71.4–5.73.54.33.51.81.2–1.71.51.32.7
Supraoccipital process-dorsal fin interdistance16.312.5–17.514.513.013.417.013.1–15.814.512.917.3
Prehyoid length17.313.1–15.414.015.614.016.716.8–19.418.114.613.9
Mouth width42.341.1–57.552.942.455.750.941.0–46.742.952.549.9
Premaxillary toothplate width9.89.9–15.912.813.514.014.211.4–15.613.713.815.0
Orbital diameter16.215.3–22.820.218.118.817.516.2–16.416.315.916.4
MeristicsMedianMedian
No.of dorsal-fin rays77–877777777
No.of anal-fin rays99–1110910109999
No.of pectoral-fin rays77–877777777
No.of caudal-fin rays1717–1817.517181515–17171717
No.of pelvic-fin rays6666666666
No.of serrae anterior margin pectoral-fin spine40–433362-53.543
No.of serrae posterior margin pectoral-fin spine148–189.5961513–15141010
Vertebrae -( -N -= 4) - -( -N -= 2) - -( -N -= 4) - -( -N -= 2) -
No.of pre-anal vertebrae1616–1716.517171514–15141414
No.of caudal vertebrae1918–201919191717–18181819
Total vertebrae3535–3635.536363232323233
- - - -Table 3. -Continued - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- - -P. luendaensis - -( -N -= 10) - - - - -P. -chiumbeensis - -( -N -= 11) - -
-Variable - -Holotype - -Range - -Mean - -Holotype - -Range - -Mean -
Standard length (mm)102.249.0–122.081.8131.553.5–142.9100.6
Body measurements (% standard length)
Preanal length59.758.9–64.261.161.459.4–63.460.8
Prepelvic length48.748.9–52.650.748.848.6–56.250.7
Predorsal length38.537.9–40.139.236.735.6–40.838.3
Prepectoral length26.826.2–29.527.726.826.7–29.527.8
Adipose fin length33.629.5–38.633.830.426.0–34.931.1
Interdorsal-adipose distance3.74.3–8.06.38.86.3–12.78.8
Dorsal fin base length16.415.8–18.217.215.715.2–17.816.1
Dorsal fin length17.017.4–24.220.618.517.3–20.919.7
Dorsal fin spine length16.714.1–19.316.816.714.5–17.415.7
Pectoral fin length18.516.4–22.620.021.519.8–23.021.0
Pectoral fin spine length16.915.2–19.017.417.416.0–19.017.6
Pelvic fin length15.114.6–19.317.213.713.6–18.816.7
Anal fin length14.212.0–15.213.613.811.8–15.713.5
Interpectoral distance18.917.3–21.118.516.915.8–20.417.4
Interpelvic distance11.54.1–10.87.09.85.4–10.87.9
Maximum body depth17.314.8–21.318.517.814.5–18.916.1
Pelvic body depth13.212.6–18.715.714.112.0–16.014.3
Minimum caudal peduncle depth12.710.9–13.011.911.19.8–12.510.9
Maximum caudal peduncle depth14.912.5–15.614.314.012.2–14.713.8
Adipose fin-caudal fin interdistance7.85.5–6.96.28.15.3–8.27.2
Anal fin-caudal fin interdistance15.112.9–15.014.415.213.6–17.615.2
Adipose fin depth6.56.4–7.76.96.25.1–6.86.1
Head length34.231.0–34.532.932.930.9–34.533.1
Head measurements (% head length)
Postorbital head length43.638.0–47.342.340.436.9–44.740.7
Preorbital head length45.144.5–48.846.247.643.4–47.746.0
Head width65.960.9–73.068.961.756.8–69.463.9
Orbital head width58.758.5–67.462.854.355.1–63.756.9
Head depth48.249.9–59.254.549.042.0–53.546.7
Orbital head depth22.931.4–37.834.134.629.6–34.132.3
Snout depth16.113.1–19.017.411.011.3–19.015.4
Maxillary barbel length57.656.1–76.165.762.248.3–68.256.8
External mandibular barbel length102.898.1–125.6111.7105.595.8–119.2105.2
- - - -Table 3. -Continued - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- -P. luendaensis -( -N -= 10) - - - -P -. -chiumbeensis -( -N -= 11) - -
-Variable -HolotypeRange -Mean -HolotypeRange -Mean -
Internal mandibular barbel length
Mandibular barbels interdistance
Interorbital distance
Anterior nostrils interdistance
Posterior nostrils interdistance
Supraoccipital process-nuchal plate interdistance
Supraoccipital process-dorsal fin interdistance
Prehyoid length
Mouth width
Premaxillary toothplate width
Orbital diameter48.040.0–56.611.310.0–15.118.817.9–23.516.810.2–19.451.912.921.214.342.536.8–49.014.310.1–13.417.616.9–21.314.112.5–16.98.27.8–11.33.71.5–5.514.014.4–22.516.014.8–18.648.241.4–48.910.78.6–13.014.315.2–19.943.411.319.314.18.28.3–10.29.39.11.21.1–5.014.214.7–23.314.410.3–17.540.940.0–52.110.810.9–13.215.414.6–19.93.118.813.045.512.117.33.419.116.545.111.317.8
Meristics
MedianMedian
No.of dorsal-fin rays77777109108–109777771717171753431412151215151717222120203736373778–107–8
No.of anal-fin rays
No.of pectoral-fin rays
No.of pelvic-fin rays
No.of caudal-fin rays17–181–59–1417–172–48–16
No.of serrae anterior margin pectoral-fin spine
No.of serrae posterior margin pectoral-fin spine
Vertebrae -( -N -= 3) -1520–2235–37 -( -N -= 4) -17–1819–2036–37
No. of pre-anal vertebrae
No.of caudal vertebrae
Total vertebrae
- - -Description: -Morphometric and meristic data are given in -Table 3 -. Body elongated. Dorsal body profile gently rising from snout tip to origin of dorsal fin, about straight from dorsal-fin origin to adipose-fin origin, and slightly directed downwards from adipose-fin origin to caudal-fin base. Body depth highest at origin of dorsal fin. Ventral body profile slightly convex from lower jaw to origin of pelvic fin, straight from pelvic fin to origin of anal fin, slightly concave from origin of anal fin to caudal-fin base. Caudal peduncle laterally compressed. Anus and urogenital opening positioned about halfway between pelvic- and anal-fin origins. - - - -Figure 6. -Photographs of the different colour patterns identified within - -Parauchenoglanis ngamensis - -. A, small-sized specimen with bars (SAIAB 204289, 36.3 mm SL, Upper Zambezi River). B, smallto medium-sized specimen with spots surrounded by light brown shading (SAIAB 204289, 46.4 mm SL, Upper Zambezi River). Scale bar: 1 cm. - - -Adipose fin longer than anal-fin base, originating anteriorly to anal-fin origin, between pelvic-fin base and anal-fin origin, and ending posteriorly to end of anal-fin base. Dorsal-fin origin anterior to pelvic-fin origin. Posterior edge of pectoral-fin rays does not reach pelvic fin. Dorsal and pectoral fins with strong spine. Posterior edge of pelvic-fin rays does not reach anal fin. Anterior margin serrated on the distal end and entire posterior margin of the pectoral-fin spine serrated. Caudal fin rounded. - -Head moderately depressed. Snout profile bluntly triangular, on dorsal view ( -Fig. 5B -). Mouth subterminal. Lips fleshy. Eyes small and situated dorsolaterally.Three pairs of barbels, base thick and tips pointed. External mandibular barbel longest, almost reaching middle of pectoral-fin spine. Inner mandibular barbel shortest, reaching middle of eye. Maxillary barbel surpassing posterior edge of eye but not reaching the base of pectoral-fin spine. Posterior nostril slits positioned about halfway between snout tip and eye. - - -Colouration in life: -Overall body colouration yellow-brown and white ventrally. Body with scattered black spots, some arranged in a series of five to seven vertical rows along the body ( -Skelton 2001: 212 -; -Marshall 2011: 146 -; - -Bruton -et al. -2018: 73 - -). - - -Colouration in alcohol: -Body brown to light brown dorsally and laterally, and light brown to white ventrally. Body with five to seven black bars in small-sized specimens, and no black spots in between bars ( -22.8–46.4 mm -SL; -Fig. 6A -). Medium-sized specimens (50.0– -97.9 mm -SL; -Fig. 6B -) with black spots smaller than the eye diameter forming five to seven vertical rows, surrounded by light brown shade. Large-sized specimens have five to seven vertical rows of black spots, smaller than eye diameter ( -101.4– 204.2 mm -SL; -Fig. 5A -). Scattered black spots present in between vertical rows in medium- and large-sized specimens. Head brown or light brown, with scattered small black spots, i.e. smaller than those on body. Fins also with numerous black spots. Distal tips of fin rays with black markings except in small-sized specimens. Usually, one or two small black spots present above gill opening and one on midline of caudal-fin base. Barbels unspotted. - - - - - -Distribution: -Parauchenoglanis ngamensis - -is known only from the -Okavango -and upper -Zambezi -(sub)basins in southern Africa ( -Fig. 1 -). - - - - -Etomologo: -Boulenger named the species after Lake Ngami, -Botswana -, where it was first recorded. - - - - \ No newline at end of file diff --git a/data/03/AE/2C/03AE2C1AA735FFB816D5FDAAFCE8FDF9.xml b/data/03/AE/2C/03AE2C1AA735FFB816D5FDAAFCE8FDF9.xml deleted file mode 100644 index d082aa68495..00000000000 --- a/data/03/AE/2C/03AE2C1AA735FFB816D5FDAAFCE8FDF9.xml +++ /dev/null @@ -1,514 +0,0 @@ - - - -Nine in one: integrative taxonomic evidence of hidden species diversity in the widespread Zambezi grunter, Parauchenoglanis ngamensis (Siluriformes: Auchenoglanididae), from southern and south-central Africa - - - -Author - -Sithole, Yonela -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Department of Ichthyology and Fisheries Science, Rhodes University, Makhanda, South Africa -y.sithole@saiab.nrf.ac.za - - - -Author - -Vreven, Emmanuel J. W. M. N. -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Vertebrate Section, Ichthyology, Royal Museum for Central Africa (RMCA), Tervuren, Belgium - - - -Author - -Bragança Tobias Musschoot, Pedro H. N. - - - -Author - -Chakona, Albert - -text - - -Zoological Journal of the Linnean Society - - -2024 - -202 - - -1 -33 - - - -journal article -10.1093/zoolinnean/zlae121 -0024-4082 -14544245 -3C2308D-7334-412D-871F-DE1F17C38D0E - - - - - - - -Parauchenoglanis chiumbeensis - -sp. nov. - - - - - - - -( -Fig. 15 -; -Table 3 -) - - - - -Parauchenoglanis ngamensis - -‘chiumbe’, - -Sithole -et al. -, 2023 - -. - - - - -Common English name: -Chiumbe grunter. - - -Common French name: -Mâchoiron de la Chiumbe. - - - - -Holotope: - - - -RMCA_Vert_P. -161724 - - -, 131.5 mm SL, -Angola -, -Kasai sub-basin -, -River Mololo -, -affluent river Chiumbe -( - -7°49 -ʹ -S - -, - -21°5 -ʹ -E - -); collector -Max Poll -; - -10 January 1963 - - -. - - -Paratopes (N = 10): - - -Angola - -: -RMCA_Vert_P.161721–161723 -, -three -, -58.2–117.5 mm -SL, collection details same as for the holotype - -. - -RMCA_Vert_P.161728−161729 -, -two -, -53.5–95.7 mm -SL, -Kasai sub-basin -, -River Kaino -( - -7°58 -ʹ -S - -, - -21°7 -ʹ -E - -); collector -Max Poll -; - -11 January 1963 - -. - - -RMCA_Vert_P.161730−161731 -, -two -, 93.8–134.0 mm SL, -Angola -, -Kasai sub-basin -, -River Tchimenji -( - -7°58 -ʹ -S - -, - -21°7 -ʹ -E - -); collector -Max Poll -; - -18 January 1963 - -. - - -SAIAB 246237 -(ex. RMCA_Vert_P.161725–161727), -three -, -93.4– 142.9 mm -SL, collection details same as for the holotype - -. - - - - - -Diagnosis: -Parauchenoglanis chiumbeensis - -is distinguished from - -P.ahli - -, - -P. altipinnis - -, - -P. balaoi - -, - -P. buetikoferi - -, - -P. longiceps - -, - -P. monkei - -, - -P. pantherinus - -, and - -P. punctatus - -by a broad humeral process (vs. pointed humeral process). It is further distinguished from - -P. altipinnis - -, - -P. balaoi - -, - -P. pantherinus - -, and - -P. punctatus - -by coarse skin (vs. smooth skin). - -Parauchenoglanis chiumbeensis - -is distinguished from - -P. zebratus - -by humeral process clearly visible through the skin and anterior margin of the pectoral-fin spine mostly smooth (vs. humeral process embedded under the skin and anterior margin of the pectoral-fin spine mostly serrated). - -Parauchenoglanis chiumbeensis - -is readily distinguished from - -P. stiassnoae - -by truncated caudal fin (vs. rounded), dorsolaterally positioned eyes (vs. dorsally positioned), and the presence of regularly spaced vertical rows of spots on the flank (vs. irregularly spaced blotches). - -Parauchenoglanis chiumbeensis - -is distinguished from - -P. ngamensis - -, - -P. lueleensis - -, - -P. poikilos - -, - -P. ernstswartzi - -, and - -P. megalasma - -by absent background spots [ -Fig. 15A -; vs. present in - -P. ngamensis - -( -Fig. 5A -), - -P. lueleensis - -( -Fig. 10A -), and - -P. poikilos - -( -Fig. 11A -) or vermiculated pattern in - -P. ernstswartzi - -( -Fig. 12A -) or blotches in - -P. megalasma - -( -Fig. 13A -)]. It is further differentiated from - -P. lueleensis - -, - -P. poikilos - -, - -P. ernstswartzi - -, and - -P. megalasma - -by relative long interdorsal–adipose distance, 6.3%– 12.7% SL (vs. narrow interdorsal–adipose distance, 3.7%–6.8% SL in - -P. lueleensis - -, 5.5%–5.8% SL in - -P. poikilos - -, 3.8%–5.0% SL in - -P. ernstswartzi - -, and 3.9%–4.8% SL in - -P.megalasma - -). It is further differentiated from - -P. ernstswartzi - -and - -P. megalasma - -by vertical rows of spots [ -Fig. 15A -; vs. vertical rows of blotches in - -P. ernstswartzi - -( -Fig. 12A -) and in - -P. megalasma - -( -Fig. 13A -)]. - -Parauchenoglanis chiumbeensis - -is differentiated from - -P. luendaensis - -by faintly spotted fins ( -Fig. 15A -; vs. unspotted in - -P. luendaensis - -, -Fig. 14A -). It is further differentiated from - -P. poikilos - -by shorter prepectoral fin length, 26.7%–29.5% SL (vs. longer, 30.6%–31.4% SL in - -P. poikilos - -). It is further differentiated from - -P. poikilos - -and - -P. megalasma - -by a shorter adipose fin, 26.0%–34.9% SL (vs. longer, 36.0%–36.6% SL in - -P. poikilos - -and 35.6%–36.9% SL in - -P. megalasma - -). It is further differentiated from - -P. dolichorhinus - -by deeper body, 14.8%–21.3% SL (vs. shallow, 11.6%–14.0% SL in - -P. dolichorhinus - -). - - - - -Description: -Morphometric and meristic data are given in -Table 3 -. Body elongated. Dorsal body profile gently rising from snout tip to origin of dorsal fin, and straight from the dorsal-fin origin to caudal-fin base. Body depth highest at origin of dorsal fin. Ventral body profile slightly convex from lower jaw to end of caudal-fin base. Caudal peduncle laterally compressed. Anus and urogenital opening positioned about halfway between pelvic- and anal-fin origins. Adipose fin longer than anal-fin base, originating anteriorly to anal-fin origin, between pelvic base and anal-fin origin, ending posteriorly to end of anal-fin base. Dorsal and pectoral fins with strong spines. Dorsal-fin origin anterior to pelvic-fin origin. Posterior tip of pectoral-fin rays does not reach pelvic fin. Posterior tip of pelvic-fin rays does not reach anal fin. Posterior margin of the pectoral-fin spine well serrated, and anterior margin mostly smooth. Caudal fin truncated. - - - -Figure 15. -Photograph of the holotype of - -Parauchenoglanis chiumbeensis -, -RMCA - -_Vert_P161724 -, 131.5 mm SL, Chiumbe River (Kasai). A, lateral view. B, dorsal view. C, ventral view. Scale bar: 1 cm. - - - -Head moderately depressed. Snout bluntly triangular, on dorsal view ( -Fig. 15B -). Mouth subterminal. Lips fleshy. Eyes small and situated dorsolaterally. Three pairs of barbels, base thick and tips pointed. External mandibular longest, reaching the distal tip of the pectoral-fin spine. Inner mandibular barbel shortest, reaching middle of the eye. Maxillary barbel reaching posterior edge of the eye. Posterior nostril slits positioned about halfway between tip of snout and eye. - - -Colouration in alcohol: -Body brown dorsally and laterally, and light brown ventrally. Body with black spots smaller than the eye forming five to seven vertical rows; however, spots surrounded by a shade on one small specimen ( -53.5 mm -SL). No additional spots in between the vertical rows, and this at all sizes. Head brown and unspotted; however, spotted on only two small specimens ( -53.5–58.2 mm -SL). All fins with faint spots. Distal tip of fin rays with no markings. Usually, two black spots present above gill opening and one spot on caudal-fin base. - - - - - -Distribution: -Parauchenoglanis chiumbeensis - -is known from the Mololo and Tchimenji rivers, left bank affluents of the Chiumbe River. - - - - - -Etomologo: -Parauchenoglanis chiumbeensis - -is named after the Chiumbe River, Kasai sub-basin, -Angola -, from which this species was collected. The suffix ‘- -ensis -’, meaning ‘lives in’, has been added. - - - - \ No newline at end of file diff --git a/data/03/AE/2C/03AE2C1AA738FFBB15A9F8EAFA58F8C0.xml b/data/03/AE/2C/03AE2C1AA738FFBB15A9F8EAFA58F8C0.xml deleted file mode 100644 index 3cad494cbc5..00000000000 --- a/data/03/AE/2C/03AE2C1AA738FFBB15A9F8EAFA58F8C0.xml +++ /dev/null @@ -1,350 +0,0 @@ - - - -Nine in one: integrative taxonomic evidence of hidden species diversity in the widespread Zambezi grunter, Parauchenoglanis ngamensis (Siluriformes: Auchenoglanididae), from southern and south-central Africa - - - -Author - -Sithole, Yonela -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Department of Ichthyology and Fisheries Science, Rhodes University, Makhanda, South Africa -y.sithole@saiab.nrf.ac.za - - - -Author - -Vreven, Emmanuel J. W. M. N. -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Vertebrate Section, Ichthyology, Royal Museum for Central Africa (RMCA), Tervuren, Belgium - - - -Author - -Bragança Tobias Musschoot, Pedro H. N. - - - -Author - -Chakona, Albert - -text - - -Zoological Journal of the Linnean Society - - -2024 - -202 - - -1 -33 - - - -journal article -10.1093/zoolinnean/zlae121 -0024-4082 -14544245 -3C2308D-7334-412D-871F-DE1F17C38D0E - - - - - - - -Parauchenoglanis megalasma - -sp. nov. - - - - - - - -( -Fig. 13 -; -Table 3 -) - - - - - -Common English name: -Large spotted grunter. - - - -Figure 13. -Photograph of the holotype of - -Parauchenoglanis megalasma -, -SAIAB - -118792 -, 72.5 mm SL, Luele River (Kasai). A, lateral view. B, dorsal view. C, ventral view. Scale bar: 1 cm. - - - -Common French name: -Mâchoiron à large taches. - - - - -Holotope: - - -SAIAB 118792 - -, -72.5 mm -SL, -Angola -, -Kasai sub-basin, -main river above confluence with tributary 1 -, -Luele -( - -7°45 -ʹ -20.1″S - -, - -19°57 -ʹ -19.4″E - -); DNA sample ES11-BT032: SB8992; collectors -Ernst Swartz -and -Fenton P.D. Cotterill -, - -16 September 2011 - - -. - - -Paratope (N = 1): - - -Angola - -: -SAIAB 235743 -(ex -SAIAB 118792 -), -one -, -66.9 mm -SL, same collection details as the holotype; DNA sample ES11-BT016: SB8993 - -. - - - - - -Diagnosis: -Parauchenoglanis megalasma - -is distinguished from - -P. ahli - -, - -P. altipinnis - -, - -P. balaoi - -, - -P. buetikoferi - -, - -P. longiceps - -, - -P. monkei - -, - -P. pantherinus - -, and - -P. punctatus - -by the presence of vertical rows of blotches (vs. vertical rows of spots or bands in - -P. monkei - -and - -P. zebratus - -). It is further differentiated from these species by a partly round snout (vs. bluntly triangular). - -Parauchenoglanis megalasma - -is distinguished from - -P. stiassnoae - -by truncated caudal fin (vs. rounded), dorsolaterally positioned eyes (vs. dorsally), and spotted fins (vs. unspotted pectoral, pelvic, and anal fins). - -Parauchenoglanis megalasma - -is distinguished from all other species except - -P -. -ernstswartzi - -by body with vertical rows of blotches with smaller blotches between them [ -Fig. 13A -; vs. body with vertical rows of spots or bars and spots between them in - -P. ngamensis - -( -Fig. 5A -) and - -P. lueleensis - -( -Fig. 10A -) or absent background spots in - -P. patersoni - -( -Fig. 11A -), - -P. dolichorhinus - -( -Fig. 8A -), - -P. luendaensis - -( -Fig. 14A -), and - -P -. -chiumbeensis - -( -Fig. 15A -)]. It is further differentiated from - -P. ernstswartzi - -by the vertical rows of blotches not extending onto adipose fin ( -Fig. 13A -; vs. rows extending onto adipose fin, -Fig. 12A -), eyes situated dorsolaterally (vs. eyes situated dorsally), and deeper body depth, 18.2%–19.9% SL (vs. shallow body depth, 15.0%–17.4% SL). - -Parauchenoglanis megalasma - -is further differentiated from - -P. luendaensis - -by spotted head and fins (vs. unspotted). It is further differentiated from - -P. luendaensis - -and - -P.chiumbeensis - -by relatively long predorsal length, 40.7%–41.9% SL (vs. shorter, 37.9%–40.1% SL in - -P. luendaensis - -and 35.6%–40.8% SL in - -P. chiumbeensis - -). It is further differentiated from - -P. chiumbeensis - -by longer adipose fin, 35.6%–36.9% SL (vs. shorter, 26.0%–34.9% SL). - - - - -Description: -Morphometric and meristic data are given in -Table 3 -. Body elongated. Dorsal body profile gently rising from snout tip to origin of dorsal fin, base of the dorsal fin slightly concave, straight from the adipose origin to caudal fin. Body depth highest at origin of dorsal fin. Ventral body profile slightly convex from lower jaw to pelvic fin origin, straight from pelvic origin to caudal fin. Caudal peduncle laterally compressed. Anus and urogenital opening positioned about halfway between pelvic- and anal-fin origins. Adipose fin longer than anal-fin base, originating anteriorly to anal-fin origin, between pelvic origin and anal-fin origin, ending posteriorly to end of anal-fin base. Dorsal and pectoral fins with strong spines. Dorsal-fin origin anterior to pelvic-fin origin. Posterior tip of pectoral-fin ray does not reach pelvic fin. Posterior margin of the pectoral-fin spine entirely serrated, and the anterior serrated towards distal end only. Caudal fin truncated. - - -Head moderately depressed. Snout profile partly round, on dorsal view ( -Fig. 13B -). Mouth subterminal. Lips fleshy. Eyes moderately large and situated dorsolaterally. Three pairs of barbels, base thick and tips pointed. External mandibular longest, almost reaching the tip of pectoral-fin spine. Inner mandibular barbel shortest, reaching posterior edge of eye. Maxillary barbel surpassing the posterior edge of the eye but not reaching the base of pectoral-fin spine. Posterior nostril slits positioned about halfway between tip of snout and eye. - - -Colouration in alcohol: -Body brown dorsally and laterally, and cream ventrally. Body with large black blotches about eye size forming six to seven vertical rows of large blotches, with those along the lateral line even bigger and thus larger than eye size. Blotches (smaller than those forming vertical rows) present in between vertical rows. Head brown with black blotches, smaller than those on body. Fins with black spots. Distal tip of fin rays with black markings. No distinct spots above gill opening, and two big spots present on caudal-fin base.Mandibular barbels (external and inner) with black spots, and maxillary barbel without spots. - - - - - -Distribution: -Parauchenoglanis megalasma - -is known from the Luele River, a right bank affluent of the Luxico River (Luchico -sensu -Poll 1967 -) and together (Luele and Luxico River) the right bank affluent of the Loange River, a left bank affluent of the -Kasai -immediately downstream of the Sankuru River, which is one of its right bank affluents. - - - - -Etomologo: -The name - -‘ -megalasma - -’ is derived from Greek ‘ -mega -’ meaning large and ‘ -melasma -’ meaning black spot, referring to the distinct large blotches along the lateral line present in this species. - - - - \ No newline at end of file diff --git a/data/03/AE/2C/03AE2C1AA739FFB41590FE18FA0FF8C3.xml b/data/03/AE/2C/03AE2C1AA739FFB41590FE18FA0FF8C3.xml deleted file mode 100644 index cff7023498d..00000000000 --- a/data/03/AE/2C/03AE2C1AA739FFB41590FE18FA0FF8C3.xml +++ /dev/null @@ -1,549 +0,0 @@ - - - -Nine in one: integrative taxonomic evidence of hidden species diversity in the widespread Zambezi grunter, Parauchenoglanis ngamensis (Siluriformes: Auchenoglanididae), from southern and south-central Africa - - - -Author - -Sithole, Yonela -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Department of Ichthyology and Fisheries Science, Rhodes University, Makhanda, South Africa -y.sithole@saiab.nrf.ac.za - - - -Author - -Vreven, Emmanuel J. W. M. N. -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Vertebrate Section, Ichthyology, Royal Museum for Central Africa (RMCA), Tervuren, Belgium - - - -Author - -Bragança Tobias Musschoot, Pedro H. N. - - - -Author - -Chakona, Albert - -text - - -Zoological Journal of the Linnean Society - - -2024 - -202 - - -1 -33 - - - -journal article -10.1093/zoolinnean/zlae121 -0024-4082 -14544245 -3C2308D-7334-412D-871F-DE1F17C38D0E - - - - - - - -Parauchenoglanis ernstswartzi - -sp. nov. - - - - - - - -( -Fig. 12 -; -Table 3 -) - - - - - -Common English name: -Ernstswartz’s grunter. - - -Common French name: -Mâchoiron d’Ernst Swartz. - - - - -Holotope: - - -SAIAB 118845 - -, -96.6 mm -SL, -Angola -, -Kasai sub-basin -, -Luxico -, -Eastern shore of Lake Carumbo -( - -7°47 -ʹ -54.6″S - -, - -19°56 -ʹ -46.8″E - -); DNA sample ES11-BT179: SB9095; collectors -Ernst Swartz -and -Fenton P.D. Cotterill -, - -4 May 2011 - - -. - - -Paratopes (N = 3): - - -Angola -: - -SAIAB 235742 -(ex -SAIAB 118845 -), -three -, 92.0– -105.6 mm -SL, locality details same as holotype, DNA sample ES11-BT178: SB9094 and ES11-BT196: SB9093 - -. - - - - - -Diagnosis: -Parauchenoglanis ernstswartzi - -is distinguished from - -P.ahli - -, - -P. altipinnis - -, - -P. balaoi - -, - -P. buetikoferi - -, - -P. longiceps - -, - -P. monkei - -, - -P. pantherinus - -, - -P. punctatus - -, and - -P. zebratus - -by eyes situated dorsally, i.e. high on the head towards its upper edge, and round snout profile (vs. eyes situated dorsally and bluntly triangular snout profile). - -Parauchenoglanis ernstswartzi - -is distinguished from - -P. stiassnoae - -by a truncated caudal fin (vs. rounded), spotted fins (vs. unspotted pectoral, pelvic and anal fin), vertical rows of blotches interspaced by reticulated pattern (vs. irregularly scattered large blotches and absence of reticulated pattern on the flank), in addition to shorter barbels, with external mandibular barbel not reaching the tip of the pectoral-fin spine (vs. barbels long, with external mandibular barbel reaching beyond the tip of the pectoral-fin spine). - -Parauchenoglanis ernstswartzi - -is distinguished from species of - -P. ngamensis - -species group by a round snout profile (vs. bluntly triangular in - -P. ngamensis - -, - -P. lueleensis - -, - -P. patersoni - -, - -P. dolichorhinus - -, - -P. poikilos - -, - -P. chiumbeensis - -, and - -P. luendaensis - -or partly round in - -P. megalasma - -); eyes situated dorsally (vs. eyes situated dorsolaterally in - -P. ngamensis - -, - -P. lueleensis - -, - -P. patersoni - -, - -P. dolichorhinus - -, - -P. poikilos - -, - -P. chiumbeensis - -, - -P. luendaensis - -, and - -P. megalasma - -), black spots at the base of the pectoral fin [ -Fig. 12C -; vs. absent in - -P. ngamensis - -( -Fig. 5C -), - -P. patersoni - -( -Fig. 11C -), - -P. dolichorhinus - -( -Fig. 8C -), - -P. lueleensis - -( -Fig. 10C -), - -P. poikilos - -( -Fig. 11C -), - -P. megalasma - -( -Fig. 13C -), - -P. chiumbeensis - -( -Fig. 14C -), and - -P. luendaensis - -( -Fig. 15C -)], black blotches (eye size) forming six to seven vertical rows extending onto adipose fin [ -Fig. 12A -; vs. five to seven vertical rows of black spots or blotches not extending onto adipose fin in - -P. ngamensis - -( -Fig. 5A -), - -P. patersoni - -( -Fig. 11A -), - -P. dolichorhinus - -( -Fig. 8A -), - -P. lueleensis - -( -Fig. 10A -), - -P. poikilos - -( -Fig. 11A -), - -P. megalasma - -( -Fig. 13A -), - -P. chiumbeensis - -( -Fig. 14A -), and - -P. luendaensis - -( -Fig. 15A -)], and vermiculated pattern on the background [ -Fig. 12A -; vs. spots on the background in - -P.ngamensis - -( -Fig. 5A -), - -P. lueleensis - -( -Fig. 10A -), and - -P. poikilos - -( -Fig. 11A -) or no background spots in - -P. patersoni - -( -Fig. 11A -), - -P. dolichorhinus - -( -Fig. 8A -), - -P. chiumbeensis - -( -Fig. 14A -), and - -P. luendaensis - -( -Fig. 15A -)]. It is further distinguished from - -P. ngamensis - -, - -P. patersoni - -, - -P. dolichorhinus - -, - -P. lueleensis - -, - -P. luendaensis - -, and - -P. chiumbeensis - -by heavily spotted barbels (vs. unspotted). - -Parauchenoglanis ernstswartzi - -is further distinguished from - -P. dolichorhinus - -by wider interpectoral distance, 19.8%–20.6% SL (vs. narrow, 12.4%–18.2% SL in - -P. dolichorhinus - -). It is further distinguished from - -P. dolichorhinus - -, - -P. lueleensis - -, and - -P. poikilos - -by deeper body depth, 18.2%–19.9% SL (vs. shallow, 11.6%–14.0% SL in - -P.dolichorhinus - -, 15.0%–17.4% SL in - -P. lueleensis - -, and 16.0%–16.2% SL in - -P. poikilos - -). - - - - -Figure 11. -Photograph of the holotype of - -Parauchenoglanis poikilos -, -SAIAB - -99100 -, 57.2 mm SL, Lovua River (Kasai). A, lateral view. B, dorsal view. C, ventral view. Scale bar: 1 cm. - - - - -Figure 12. -Photograph of the holotype of - -Parauchenoglanis ernstswartzi -, -SAIAB - -118845 -, 96.6 mm SL, Lake Carumbo (Kasai). A, lateral view. B, dorsal view. C, ventral view. Scale bar: 1 cm. - - - - -Description: -Morphometric and meristic data are given in -Table 3 -. Body elongated. Dorsal body profile flat, slightly rising from snout tip to origin of dorsal fin, straight from dorsal-fin origin to caudal-fin base. Body depth highest at origin of dorsal fin. Ventral body profile convex from lower jaw to end of caudal-fin base. Caudal peduncle laterally compressed. Anus and urogenital opening positioned about halfway between pelvic- and anal-fin origins. Adipose fin longer than anal-fin base, originating at the base of dorsal fin, ending posteriorly to end of anal-fin base. Dorsal and pectoral fins with strong spines. Dorsal-fin short with long base, originating anterior to pelvic-fin origin. Pectoral fin with strong spine. Posterior tip of pectoral-fin rays does not reach pelvic fins. Posterior tip of pelvic-fin rays does not reach anal fin. Entire posterior margin of the pectoral-fin spine serrated, and anterior margin serrated on the distal end. Caudal fin truncated. Fleshy flanges along the dorsal and ventral edges of the base of caudal fin. - - -Head moderately depressed. Snout profile round, on dorsal view ( -Fig. 12B -). Mouth subterminal. Lips fleshy. Eyes big and situated dorsally, orbits projecting on the head. Three pairs of barbels, base thick and tips pointed. External mandibular barbel longest, almost reaching the tip of pectoral-fin spine. Inner mandibular barbel shortest, reaching posterior edge of eye. Maxillary barbel reaching behind posterior edge of the eye but not reaching the base of pectoral-fin spine. Posterior nostril slits positioned about halfway between snout tip and eye. - - -Colouration in alcohol: -Body black dorsally, light brown laterally (below lateral line), and cream ventrally. Body with black blotches (i.e. irregularly shaped spots) about eye size, forming six to seven vertical rows, with rows also extending onto adipose fin. Vermiculated pattern present in between vertical rows. Head black, with black blotches smaller than those on body. Fins with black spots arranged along the rays. Distal tip of fin rays with black markings. Barbels with numerous black spots. - - - - - -Distribution: -Parauchenoglanis ernstswartzi - -is currently reported from the Lake Carumbo on the Luxico River, the upstream part of the Loange River, a left bank affluent of the -Kasai -River (see -Poll 1967 -), the latter being a left bank affluent of the -middle Congo -Basin. - - - - -Etomologo: -The species ephitet - -‘ -ernstswartzi - -’ is dedicated to Dr Ernst Swartz, who collected specimens used in the description of six new species in this study. This dedication is in recognition of his pioneering and extensive exploration of the Kwanza and adjacent river systems, including the upper -Kasai -sub-basin. - - - - \ No newline at end of file diff --git a/data/03/AE/2C/03AE2C1AA73AFFB5159EFB80FB06FEB2.xml b/data/03/AE/2C/03AE2C1AA73AFFB5159EFB80FB06FEB2.xml deleted file mode 100644 index 691ab688625..00000000000 --- a/data/03/AE/2C/03AE2C1AA73AFFB5159EFB80FB06FEB2.xml +++ /dev/null @@ -1,383 +0,0 @@ - - - -Nine in one: integrative taxonomic evidence of hidden species diversity in the widespread Zambezi grunter, Parauchenoglanis ngamensis (Siluriformes: Auchenoglanididae), from southern and south-central Africa - - - -Author - -Sithole, Yonela -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Department of Ichthyology and Fisheries Science, Rhodes University, Makhanda, South Africa -y.sithole@saiab.nrf.ac.za - - - -Author - -Vreven, Emmanuel J. W. M. N. -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Vertebrate Section, Ichthyology, Royal Museum for Central Africa (RMCA), Tervuren, Belgium - - - -Author - -Bragança Tobias Musschoot, Pedro H. N. - - - -Author - -Chakona, Albert - -text - - -Zoological Journal of the Linnean Society - - -2024 - -202 - - -1 -33 - - - -journal article -10.1093/zoolinnean/zlae121 -0024-4082 -14544245 -3C2308D-7334-412D-871F-DE1F17C38D0E - - - - - - - -Parauchenoglanis poikilos - -sp. nov. - - - - - - - -( -Fig. 11 -; -Table 3 -) - - - - - -Common English name: -Spotted grunter. - - -Common French name: -Mâchoiron tachetés. - - - - -Holotope: - - -SAIAB 99100 - -, -57.2 mm -SL, -Angola -, -Kasai sub-basin -, -Lovua -, -Opposite Capeia Village on bamboo track -( - -8°20 -ʹ -16.3″S - -, - -20°14 -ʹ -29.0″E - -); DNA sample ES11-BT218: SB9000; collectors -Ernst Swartz -and -Fenton P.D. Cotterill -; - -5 July 2011 - - -. - - -Paratope (N = 1): - - -Angola - -: -SAIAB 235744 -(ex -SAIAB 99100 -), -one -, -36.3 mm -SL, collection details same as for holotype; DNA sample ES11-BT204: SB8977 - -. - - - - - -Diagnosis: -Parauchenoglanis poikilos - -is distinguished from - -P. ahli - -, - -P. altipinnis - -, - -P. balaoi - -, - -P. buetikoferi - -, - -P. longiceps - -, - -P. monkei - -, - -P. pantherinus - -, and - -P. punctatus - -by a broad humeral process (vs. pointed humeral process). It is further distinguished from - -P. altipinnis - -, - -P. balaoi - -, - -P. pantherinus - -, and - -P. punctatus - -by coarse skin (vs. smooth skin). It is differentiated from - -P. buetikoferi - -and - -P. monkei - -by anterior margin of the pectoral-fin spine mostly smooth (vs. anterior margin of pectoral-fin spine serrated from base to tip). - -Parauchenoglanis poikilos - -is distinguished from - -P. zebratus - -by humeral process clearly visible through the skin and anterior margin of the pectoral-fin spine mostly smooth (vs. humeral process embedded under the skin and anterior margin of the pectoral-fin spine mostly serrated). - -Parauchenoglanis poikilos - -is distinguished from - -P. stiassnoae - -by dorsolaterally positioned eyes (vs. dorsally) and spotted fins (vs. unspotted pectoral, pelvic and anal fin). - -Parauchenoglanis poikilos - -is distinguished from - -P. patersoni - -, - -P. dolichorhinus - -, - -P. luendaensis - -, and - -P. chiumbeensis - -by presence of black background spots in between the darker spots constituting the vertical rows ( -Fig. 11A -; vs. absent, -Figs 11A -, -13A -, -14A -, -15A -, respectively). It is further distinguished from - -P -. -luendaensis - -by spotted head and fins ( -Fig. 11B -; vs. unspotted, -Fig. 14B -). It is distinguished from - -P -. -ernstswartzi - -and - -P -. -megalasma - -by body with vertical rows of spots ( -Fig. 11A -; vs. vertical rows of blotches, -Fig. 13A -) and bluntly triangular snout [ -Fig. 11B -; vs. round snout in - -P. ernstswartzi - -( -Fig. 12B -) or partly round snout in - -P. megalasma - -( -Fig. 13B -)]. - -Parauchenoglanis poikilos - -is further distinguished from - -P. ernstswartzi - -by lower body depth, 16.0%– 16.2% SL (vs. deeper, 18.2%–19.9% SL in - -P. ernstswartzi - -). It is distinguished from - -P. dolichorhinus - -by deeper body, 16.0%– 16.2% SL (vs. shallow, 11.6%–14.0% SL in - -P. dolichorhinus - -). It is differentiated from - -P. lueleensis - -by scattered black spots in between vertical rows ( -Fig. 11A -; vs. one or two spots in between vertical rows, -Fig. 10A -) and spotted mandibular barbels ( -Fig. 10B -; vs. unspotted, -Fig. 10B -). - - - - -Description: -Morphometric and meristic data are given in -Table 3 -. Body elongated. Dorsal body profile gently rising from snout tip to origin of dorsal fin, slightly concave from dorsal-fin origin to adipose-fin origin, and straight from the adipose-fin origin to caudal-fin base. Body depth highest at origin of dorsal fin. Ventral body profile slightly convex from lower jaw to end of caudal-fin base. Caudal peduncle laterally compressed. Anus and urogenital opening positioned about halfway between pelvic- and anal-fin origins. Adipose fin longer than anal-fin base, originating anteriorly to anal-fin origin, between pelvic origin and anal-fin origin, ending posteriorly to end of anal-fin base. Dorsal and pectoral fins with strong spines. Dorsal-fin origin anterior to pelvic-fin origin. Posterior edge of pectoral-fin rays does not reach pelvic fin. Pectoral fine spine serrated along inner posterior margin and towards extremity of outer anterior margin. Caudal fin truncated. - - -Head moderately depressed. Snout profile bluntly triangular, on dorsal view ( -Fig. 11B -). Mouth subterminal. Lips fleshy. Eyes small and situated dorsolaterally. Three pairs of barbels, base thick and tips pointed. External mandibular barbel longest, almost reaching the tip of pectoral-fin spine. Inner mandibular barbel shortest, reaching posterior edge of the eye. Maxillary barbel reaching behind posterior edge of the eye but not reaching the base of pectoral-fin spine. Posterior nostril slits positioned about halfway between tip of snout and eye, and both closer together than the interorbital distance. - - -Colouration in alcohol: -Body brown dorsally and laterally, and cream ventrally. Body with black spots smaller than the eye surrounded by a light brown shade forming six to seven vertical rows. Scattered black spots present in between vertical rows. Head brown, with numerous small black spots, smaller than those on body. Fins with black spots arranged along the rays. No markings on distal tips of fin rays. Two large brown blotches on the end of the caudal peduncle surrounded by a light yellowish brown area and delimited posteriorly by a crescent-shaped marking present on the caudal-fin base. Mandibular barbels (external and inner) with black spots, and maxillary barbel without spots. - - - - - -Distribution: -Parauchenoglanis poikilos - -is currently known from the Lovua River (Luchico -sensu -Poll 1967 -), a left bank affluent of the -Kasai -River (see -Poll 1967 -), with the latter being a left bank affluent of the -middle Congo -Basin. - - - - -Etomologo: -The name - -‘ -poikilos - -’ is derived from the Greek adjective - -‘ -poikilos - -’, meaning spotted, in reference to the numerous spots composing its colour pattern. - - - - \ No newline at end of file diff --git a/data/03/AE/2C/03AE2C1AA73BFFB61587FE9DFCE8FC29.xml b/data/03/AE/2C/03AE2C1AA73BFFB61587FE9DFCE8FC29.xml deleted file mode 100644 index 082d80d697f..00000000000 --- a/data/03/AE/2C/03AE2C1AA73BFFB61587FE9DFCE8FC29.xml +++ /dev/null @@ -1,549 +0,0 @@ - - - -Nine in one: integrative taxonomic evidence of hidden species diversity in the widespread Zambezi grunter, Parauchenoglanis ngamensis (Siluriformes: Auchenoglanididae), from southern and south-central Africa - - - -Author - -Sithole, Yonela -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Department of Ichthyology and Fisheries Science, Rhodes University, Makhanda, South Africa -y.sithole@saiab.nrf.ac.za - - - -Author - -Vreven, Emmanuel J. W. M. N. -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Vertebrate Section, Ichthyology, Royal Museum for Central Africa (RMCA), Tervuren, Belgium - - - -Author - -Bragança Tobias Musschoot, Pedro H. N. - - - -Author - -Chakona, Albert - -text - - -Zoological Journal of the Linnean Society - - -2024 - -202 - - -1 -33 - - - -journal article -10.1093/zoolinnean/zlae121 -0024-4082 -14544245 -3C2308D-7334-412D-871F-DE1F17C38D0E - - - - - - - -Parauchenoglanis lueleensis - -sp. nov. - - - - - - - -( -Fig. 10 -; -Table 3 -) - - - - - -Common English name: -Luele grunter. - - -Common French name: -Mâchoiron de la Luele. - - - - -Holotope: - - -SAIAB 118796 - -, -95.4 mm -SL, -Angola -, -Kasai sub-basin -, -Luxico -, -above drift in tributary 2 -( - -7°45 -ʹ -05.7″S - -, - -19°57 -ʹ -18.8″E - -); DNA sample ES11-BT103: SB8998; collectors -Ernst Swartz -and -Cotterill P.D. Fenton -; - -29 April 2011 - - -. - - -Paratopes (N = 13): - - -Angola - -: -RMCA_Vert_P.0003 -(ex SAIAB 118850), -one -, -158.5 mm -SL, -Kasai sub-basin -, -Luxico -, above -confluence with main river in tributary 2 -( - -7°45 -ʹ -7.0″S - -, - -19°57 -ʹ -19.6″E - -); DNA sample ES11-BT049: SB8999; collectors -Ernst Swartz -and -Cotterill P.D. Fenton -; - -29 April 2011 - - -. - -SAIAB 99425 -, -three -, -53.2–91.6 mm -SL, -Kasai sub-basin -, -Luxico -, -above confluence with main river in tributary -1 ( - -7°45 -ʹ -09.2″S - -, - -19°57 -ʹ -22.8″E - -); DNA sample ES11-BT124: SB8997; collectors -Ernst Swartz -and -Cotterill P.D. Fenton -; - -2 May 2011 - - -. - -SAIAB 99230 -, five, 39.0– -21.8 mm -SL, -Kasai sub-basin -, -Luxico -, -Luele Rapids above the confluence with the Luxico River -( - -7°45 -ʹ -01.6″S - -, - -19°56 -ʹ -58.8″E - -); DNA sample ES11-BT083: SB8990 and ES11-BT084: SB8991; collectors -Ernst Swartz -and -Cotterill P.D. Fenton -; - -1 May 2011 - - -. - -SAIAB 99421 -, -four -, -17.2–70.72mm -SL, -Kasai sub-basin -, -Luxico -, -between Lake Carumbo and the confluence with the Luele -River ( - -7°45 -ʹ -29.4″S - -, - -19°56 -ʹ -34.3″E - -); DNA sample ES11-BT060: SB8994 and ES11-BT059: SB8995; collectors -Ernst Swartz -and -Cotterill P.D. Fenton -; - -1 May 2011 - - -. - - - - -Figure 9. -Photograph of the typical colour pattern of a small-sized specimen of - -Parauchenoglanis dolichorhinus -, SAIAB - -85066, - - - -Figure 10. -Photograph of the holotype of - -Parauchenoglanis lueleensis -, -SAIAB - -118796 -, 95.4 mm SL, Luele River (Kasai). A, lateral view. B, dorsal view. C, ventral view. Scale bar: 1 cm. - - -52.4 mm SL, Kwanza River. Scale bar: 1 cm. - - - - -Diagnosis: -Parauchenoglanis lueleensis - -is distinguished from - -P. ahli - -, - -P. altipinnis - -, - -P. balaoi - -, - -P. buetikoferi - -, - -P. longiceps - -, - -P. monkei - -, - -P. pantherinus - -, and - -P. punctatus - -by a broad humeral process (vs. pointed humeral process). It is further distinguished from - -P. altipinnis - -, - -P. balaoi - -, - -P. pantherinus - -, and - -P. punctatus - -by coarse skin (vs. smooth skin). It is differentiated from - -P. buetikoferi - -and - -P. monkei - -by anterior margin of the pectoral-fin spine mostly smooth (vs. anterior margin of pectoral-fin spine serrated from base to tip). - -Parauchenoglanis lueleensis - -is distinguished from - -P. zebratus - -by humeral process clearly visible through the skin, presence of six to seven vertical rows of spots regardless of the size, and anterior margin of the pectoral-fin spine mostly smooth (vs. humeral process embedded under the skin, four to five vertical rows of spots on small specimens or four to five bars on larger specimens, and anterior margin of the pectoral-fin spine mostly serrated). - -Parauchenoglanis lueleensis - -is distinguished from - -P. stiassnoae - -by the dorsolaterally positioned eyes (vs. dorsally) and spotted fins (vs. unspotted pectoral, pelvic, and anal fin). - -Parauchenoglanis lueleensis - -is distinguished from all other species in the - -P. ngamensis - -group by one or two black background spots in between vertical rows of spots [ -Fig. 10A -; vs. absent background spots in - -P. patersoni - -( -Fig. 11A -), - -P -. -dolichorhinus - -( -Fig. 8A -), - -P -. -luendaensis - -( -Fig. 14A -), and - -P -. -chiumbeensis - -( -Fig. 15A -) or numerous spots in - -P. ngamensis - -( -Fig. 5A -) and - -P -. -poikilos - -( -Fig. 11A -) or vermiculated pattern on the background in - -P -. -ernstswartzi - -( -Fig. 12A -) or blotches in - -P -. -megalasma - -( -Fig. 13A -)]. It is further differentiated from - -P -. -luendaensis - -by spotted fins ( -Fig. 10A -; vs. unspotted in - -P -. -luendaensis - -, -Fig. 14A -). It is further differentiated from - -P -. -ernstswartzi - -and - -P -. -megalasma - -by shallow body depth, 15.0%–17.4% SL (vs. deeper, 18.2%–19.9% SL in - -P. ernstswartzi - -and 20.2%–20.9% SL in - -P -. -megalasma - -). It is further differentiated from - -P. dolichorhinus - -by deeper head, 41.2%–50.3% SL (vs. smaller head depth in - -P. dolichorhinus - -, 34.6%–40.0% SL) and wider interpectoral distance, 18.8%–22.5% SL (vs. narrow interpectoral distance in - -P -. -dolichorhinus - -, 17.4%–18.2% SL). - - - - -Description: -Morphometric and meristic data are given in -Table 3 -. Body elongated. Dorsal body profile gently rising from snout tip to dorsal-fin origin, concave from dorsal-fin origin to adipose-fin origin, and straight from the adipose-fin origin to caudal-fin base. Body depth highest at origin of dorsal fin. Ventral body profile slightly convex from lower jaw to end of caudal-fin base. Caudal peduncle laterally compressed. Anus and urogenital opening positioned about halfway between pelvic- and anal-fin origins. - -Adipose fin longer than anal-fin base, originating anteriorly to anal-fin origin, between pelvic base and anal-fin origin, ending posteriorly to end of anal-fin base. Dorsal and pectoral fins with strong spine. Dorsal-fin origin anterior to pelvic-fin origin. Posterior tip of pectoral-fin rays does not reach pelvic fin. Entire posterior margin of the pectoral-fin spine serrated, and anterior serrated only towards distal end. Caudal fin truncated. - -Head moderately depressed. Snout profile bluntly triangular, on dorsal view ( -Fig. 10B -). Mouth subterminal. Lips fleshy. Eyes small and situated dorsolaterally. Three pairs of barbels, base thick and tips pointed. External mandibular barbel longest, almost reaching the tip of pectoral-fin spine. Inner mandibular barbel shortest, reaching middle of eye. Maxillary barbel reaching beyond posterior edge of the eye but not reaching the base of pectoral-fin spine. Posterior nostril slits positioned about halfway between snout tip and eye. - - -Colourationinalcohol: -Bodybrowndorsallyandlaterally,andcream ventrally. Body with black spots smaller than the eye forming six to seven vertical rows of black spots, at all sizes. One or two faint spots present in between vertical rows. Head brown, with black spots, smaller than those on body mostly in small-sized specimens and sometimes even unspotted or with small black spots present only on edge of the operculum. Fins with small black spots. No markings on distal tips of fin rays. Usually, one or two small black spots present above the gill opening. Barbels unspotted. - - - - - -Distribution: -Parauchenoglanis lueleensis - -is known from the Luele River and its right bank affluent, Luxico River (Luchico -sensu -Poll 1967 -). Both are tributaries of the Loange River, a left bank affluent of the -Kasai -River (see -Poll 1967 -), itself a left bank affluent of the -middle Congo -Basin. - - - - - -Etomologo: -Parauchenoglanis lueleensis - -is named after the Luele River, Kasai sub-basin, -Angola -, where this species is currently known to occur. The suffix ‘- -ensis -’, meaning ‘lives in’, has been added. - - - - \ No newline at end of file diff --git a/data/03/AE/2C/03AE2C1AA73CFFB716E0FCEEFACDFEC9.xml b/data/03/AE/2C/03AE2C1AA73CFFB716E0FCEEFACDFEC9.xml deleted file mode 100644 index 747f329845f..00000000000 --- a/data/03/AE/2C/03AE2C1AA73CFFB716E0FCEEFACDFEC9.xml +++ /dev/null @@ -1,528 +0,0 @@ - - - -Nine in one: integrative taxonomic evidence of hidden species diversity in the widespread Zambezi grunter, Parauchenoglanis ngamensis (Siluriformes: Auchenoglanididae), from southern and south-central Africa - - - -Author - -Sithole, Yonela -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Department of Ichthyology and Fisheries Science, Rhodes University, Makhanda, South Africa -y.sithole@saiab.nrf.ac.za - - - -Author - -Vreven, Emmanuel J. W. M. N. -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Vertebrate Section, Ichthyology, Royal Museum for Central Africa (RMCA), Tervuren, Belgium - - - -Author - -Bragança Tobias Musschoot, Pedro H. N. - - - -Author - -Chakona, Albert - -text - - -Zoological Journal of the Linnean Society - - -2024 - -202 - - -1 -33 - - - -journal article -10.1093/zoolinnean/zlae121 -0024-4082 -14544245 -3C2308D-7334-412D-871F-DE1F17C38D0E - - - - - - - -Parauchenoglanis dolichorhinus - -sp. nov. - - - - - - - -( -Fig. 8 -) - - - - - -Common English name: -Long-snouted grunter. - - -Common French name: -Mâchoiron à long museau. - - - - -Holotope: - - - -SAIAB -84886 - - -, 139.8 mm SL, -Angola -, -Kwanza Basin -, -Posto 5 -( - -09°48 -ʹ -23″S - -, - -15°24 -ʹ -30″E - -); DNA sample ES07F442: SB9092; collector -Ernst Swartz -, - -31 October 2007 - - -. - - -Paratopes (N = 3): - - -Angola - -: -RMCA_Vert_P.0002 -(ex SAIAB 235745), -one -, -115.1 mm -SL, -Kwanza Basin -, -Posto 5 -( - -09°48 -ʹ -23″S - -, - -15°24 -ʹ -30″E - -); DNA sample ES07F296: SB9090; collector -Ernst Swartz -, - -24 October 2007 - - -. - -SAIAB 235745 -, -one -, -91.9 mm -SL, -Kwanza Basin -, -Posto 5 -( - -09°48 -ʹ -23″S - -, - -15°24 -ʹ -30″E - -); collector -Ernst Swartz -, - -24 October 2007 - - -. - -SAIAB 85066 -, -one -, -52.4 mm -SL, -Kwanza Basin -, -New dam site -( - -09°41 -ʹ -35″S - -, - -14°59 -ʹ -55″E - -); DNA sample ES08B148: -SAFW418-08 -; collector -Ernst Swartz -, - -16 August 2008 - - -. - - - - - -Diagnosis: -Parauchenoglanis dolichorhinus - -is distinguished from - -P.ahli - -, - -P. altipinnis - -, - -P. balaoi - -, - -P. buetikoferi - -, - -P. longiceps - -, - -P. monkei - -, - -P. pantherinus - -, and - -P. punctatus - -by a broad humeral process (vs. pointed humeral process). It is further distinguished from - -P. altipinnis - -, - -P. balaoi - -, - -P. pantherinus - -, and - -P. punctatus - -by coarse skin (vs. smooth skin). It is differentiated from - -P. buetikoferi - -and - -P. monkei - -by anterior margin of the pectoral-fin spine mostly smooth (vs. anterior margin of pectoral-fin spine serrated from base to tip). - -Parauchenoglanis dolichorhinus - -is distinguished from - -P. zebratus - -by humeral process clearly visible through the skin, presence of five to six bars on small or vertical rows of spots on large, and anterior margin of the pectoral-fin spine mostly smooth (vs. humeral process embedded under the skin, four to five vertical rows of spots on small specimens or four to five bars on larger specimens, and anterior margin of the pectoral-fin spine mostly serrated). - -Parauchenoglanis dolichorhinus - -is distinguished from - -P. stiassnoae - -by the absence of black background spots in between the vertical bars or rows of spots (vs. reticulated pattern), a depressed head, head depth 34.6%–40.0% HL (vs. moderately depressed head, head depth 43.5%–58.7% HL), and a narrow body depth, 11.6%–14.0% SL (vs. a deeper body depth, 15.7%–22.7% SL). - -Parauchenoglanis dolichorhinus - -is distinguished from all other species of the - -P. ngamensis - -species group by a depressed head; head depth 34.6%–40.0% HL (vs. a moderately depressed head; head depth 40.6%–60.8% HL in - -P. ngamensis - -, 48.5%–54.9% HL in - -P. patersoni - -, 41.2%–50.3% HL in - -P. lueleensis - -, 41.7%–42.0% HL in - -P. poikilos - -, 49.5%–54.0% HL in - -P. ernstswartzi - -, 42.0%–53.5% HL in - -P. chiumbeensis - -, and 48.2%– 59.2% HL in - -P.luendaensis - -). It is further distinguished from these species by narrow body depth, 11.6%–14.0% SL (vs. deeper body depth, 14.6%–23.5% SL in - -P. ngamensis - -, 15.5%–21.4% SL in - -P. patersoni - -, 15.0%–17.4% SL in - -P. lueleensis - -, 16.0%–16.2% SL in - -P. poikilos - -, 18.2%–19.9% SL in - -P. ernstswartzi - -, 20.2%–20.9% SL in - -P. megalasma - -, 14.8%–21.3% SL in - -P. luendaensis - -, and 14.5%–18.9% SL in - -P. chiumbeensis - -). It is further distinguished from - -P. lueleensis - -, - -P. poikilos - -, - -P. ernstswartzi - -, and - -P. megalasma - -by a narrow interpectoral distance, 17.4%–18.2% SL (vs. wider, 18.8%–22.5% SL in - -P. lueleensis - -, 18.4%–21.0% SL in - -P. poikilos - -, 19.8%–20.6% SL in - -P. ernstswartzi - -, and 18.8%–21.9 % SL in - -P. megalasma - -). Finally, it is further distinguished by unspotted fins [ -Fig. 8A -; vs. fins with numerous black spots in - -P. ngamensis - -( -Fig. 5A -), - -P. lueleensis - -( -Fig. 10A -), and - -P. poikilos - -( -Fig. 11A -) or faint spots in - -P. chiumbeensis - -( -Fig. 15A -) or blotches in - -P. ernstswartzi - -( -Fig. 12A -) and - -P. megalasma - -( -Fig. 13A -)]. - - - - -Description: -Morphometric and meristic data are given in -Table 3. A -medium-sized species with a maximum observed size of -139.8 mm -SL. Body elongated and dorsally depressed from snout tip to origin of dorsal fin. Dorsal body profile convex from tip of snout to dorsal fin origin, straight from dorsal-fin origin to caudal-fin base. Body depth highest at origin of dorsal fin. Ventral body profile slightly concave from lower jaw to caudal-fin base. Caudal peduncle laterally compressed. Anus and urogenital opening positioned about halfway between pelvic- and anal-fin origins. - - - -Figure 8. -Photograph of the holotype of - -Parauchenoglanis dolichorhinus -, -SAIAB - -84886 -, 139.8 mm SL, Kwanza River. A, lateral view. B, dorsal view. C, ventral view. Scale bar: 1 cm. - - -Adipose fin longer than anal-fin base, originating anteriorly to anal-fin origin, between pelvic base and anal-fin origin, ending posteriorly to end of anal-fin base. Dorsal and pectoral fins with strong spine. Dorsal-fin origin anterior to pelvic-fin origin. Posterior edge of pectoral fin rays does not reach pelvic fins. Entire posterior margin of the pectoral-fin spine serrated, and anterior margin serrated only towards distal end. Caudal fin rounded. - -Head depressed. Snout profile bluntly triangular, on dorsal view ( -Fig. 8B -). Mouth subterminal. Lips fleshy. Eyes small and situated dorsolaterally. Three pairs of barbels, base thick and tips pointed. External mandibular barbel longest, almost reaching the tip of pectoral-fin spine. Inner mandibular barbel shortest, reaching middle of eye. Maxillary barbel reaching beyond posterior edge of the eye but not reaching the base of pectoral-fin spine. Posterior nostril slits positioned about halfway between snout tip and eye. - - -Colouration in life: -Body yellow-brown to light brown dorsally and laterally. Distal tips of fins with brown markings (Supporting Information, -Fig. S3 -). Arrangement of spots is discussed below. - - -Colouration in alcohol: -Body light brown dorsally and laterally, and cream ventrally. Body with five to six bars in medium-sized specimens ( -52.4–91.9 mm -SL; -Fig. 9 -). Large-sized specimens ( -115.1–139.8 mm -SL) with black spots smaller than the eye forming six vertical rows. No additional black spots in between the bars or vertical rows of spots. Head light brown, sometimes with very few, usually <10 black spots, smaller than those on body. Fins with a few black spots, sometimes even unspotted. Distal tip of fin rays with black markings. Usually, two small black spots present above gill opening and one spot on midline of caudal-fin base. Barbels unspotted. - - - - - -Distribution: -Parauchenoglanis dolichorhinus - -has currently been recorded only from the mainstream section of the Kwanza Basin above and below the Quissaquina Falls ( -Fig. 1 -). This species co-occurs with the other Kwanza endemic - -Parauchenoglanis -species - -, - -P. patersoni - -. - - - - -Etomologo: -The species name - -‘ -dolichorhinus - -’ is derived from the Greek ‘ -dolichos -’ meaning long and ‘ -rhinus -’ meaning snout, referring to the long snout (preorbital length) of this species in comparison to the other species in the - -P. ngamensis - -group. - - - - \ No newline at end of file diff --git a/data/03/AE/2C/03AE2C1AA73EFFB01596FB61FDFEFCDF.xml b/data/03/AE/2C/03AE2C1AA73EFFB01596FB61FDFEFCDF.xml deleted file mode 100644 index e05b646bd0c..00000000000 --- a/data/03/AE/2C/03AE2C1AA73EFFB01596FB61FDFEFCDF.xml +++ /dev/null @@ -1,589 +0,0 @@ - - - -Nine in one: integrative taxonomic evidence of hidden species diversity in the widespread Zambezi grunter, Parauchenoglanis ngamensis (Siluriformes: Auchenoglanididae), from southern and south-central Africa - - - -Author - -Sithole, Yonela -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Department of Ichthyology and Fisheries Science, Rhodes University, Makhanda, South Africa -y.sithole@saiab.nrf.ac.za - - - -Author - -Vreven, Emmanuel J. W. M. N. -National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda, South Africa & Vertebrate Section, Ichthyology, Royal Museum for Central Africa (RMCA), Tervuren, Belgium - - - -Author - -Bragança Tobias Musschoot, Pedro H. N. - - - -Author - -Chakona, Albert - -text - - -Zoological Journal of the Linnean Society - - -2024 - -202 - - -1 -33 - - - -journal article -10.1093/zoolinnean/zlae121 -0024-4082 -14544245 -3C2308D-7334-412D-871F-DE1F17C38D0E - - - - - - - -Parauchenoglanis patersoni - -sp. nov. - - - - - - - -( -Fig. 7 -; -Table 3 -) - - - - - -Common English name: -Paterson’s grunter. - - -Common French name: -Mâchoiron de Paterson. - - - - -Holotope: - - - -SAIAB -84804 - - -, 192.9 mm SL, -Angola -, -Kwanza Basin -, -Posto 5 -( - -9°48 -ʹ -23″S - -, - -15°24 -ʹ -30″E - -); DNA sample ES07F293: SB9098; collector -Ernst Swartz -, - -24 October 2007 - - -. - - -Paratopes (N = 14): - - -Angola - -: -RMCA_Vert_P.0001 -(ex -SAIAB 235741 -), -one -, 215.0 mm SL, collection details same as for the holotype; -Kwanza -, -Posto 5 -( - -9°48 -ʹ -23″S - -, - -15°24 -ʹ -30″E - -); DNA sample ES07F295: SB9089; collector -Ernst Swartz -, - -24 October 2007 - - -. - -SAIAB 235741 -(ex -SAIAB 84804 -), -two -, 185.1–206.0 mm SL, collection details same as for the holotype; -Kwanza -, -Posto 5 -( - -9°48 -ʹ -23″S - -, - -15°24 -ʹ -30″E - -); collector -Ernst Swartz -, - -24 October 2007 - -. - - -SAIAB 84844 -, -one -, 223.0 mm SL, -Kwanza River -, -Second Fishing village -( - -9°46 -ʹ -21″S - -, - -15°32 -ʹ -31″E - -), DNA sample ES07F374: SB9091; collector -Ernst Swartz -; - -15 September 2009 - -. - - -SAIAB 85190 -, -one -, 117.0 mm SL, -Kwanza Basin -, -Luando River -, -Bridge between Cimanga and Capunda -( - -10°38 -ʹ -26″S - -, - -17°25 -ʹ -06″E - -); DNA sample ES08B308: -SAFW563-09 -; collector -Ernst Swartz -; - -18 September 2009 - -. - - -SAIAB 235747 -(ex -SAIAB 84804 -), -two -, -52.7−65.6 mm -SL, collection details same as for the holotype; - - -SAIAB 235746 -(ex SAIAB 85190), -five -, -42.9–116.1 mm -SL, -Kwanza Basin -, -Luando River -, -Bridge between Cimanga and Capunda -( - -10°38 -ʹ -26″S - -, - -17°25 -ʹ -06″E - -); DNA sample ES08B310: SAFW565-09; collector -Ernst Swartz -; - -18 September 2009 - -. - - -SAIAB 84625, -one -, -88.8 mm -SL, -Kwanza Basin -, -Terra Nova village -above bridge at Cambambe inflow -( - -9°46 -ʹ -44.0″S - -, - -14°31 -ʹ -24.0″E - -); DNA sample ESO7F041: SAFW237-08; collector -Ernst Swartz -; - -10 October 2007 - - -. - -SAIAB 85520 -, -one -, -59.5 mm -SL, -Kwanza Basin -, -Posto 5 -( - -9°48 -ʹ -23.0″S - -, - -15°24 -ʹ -30.0″E - -); DNA sample: ES07D096: SB9097; collectors -Ernst Swartz, D. Neto -and -Paul Skelton -; - -19 August 2007 - -. - - - - - -Figure 7. -Photograph of the holotype of - -Parauchenoglanis patersoni -, -SAIAB - -84804 -, 192.9 mm SL, Kwanza River. A, lateral view. B, dorsal view. C, ventral view. Scale bar: 1 cm. - - - - - -Diagnosis: -Parauchenoglanis patersoni - -is distinguished from - -P. ahli - -, - -P. altipinnis - -, - -P. balaoi - -, - -P. buetikoferi - -, - -P. longiceps - -, - -P. monkei - -, - -P. pantherinus - -, and - -P. punctatus - -by a broad humeral process (vs. pointed humeral process). It is further distinguished from - -P. altipinnis - -, - -P. balaoi - -, - -P. pantherinus - -, and - -P. punctatus - -by a coarse skin (vs. smooth skin). It is differentiated from - -P. buetikoferi - -and - -P. monkei - -by anterior margin of the pectoral-fin spine mostly smooth (vs. anterior margin of pectoral-fin spine serrated from base to tip). - -Parauchenoglanis patersoni - -is distinguished from - -P. zebratus - -by a humeral process clearly visible through the skin, presence of five to six bars on small specimens or vertical rows of spots on large specimens (rarely seven), and anterior margin of the pectoral-fin spine mostly smooth (vs. humeral process embedded under the skin, four to five vertical rows of spots on small or four to five bars on large specimens, and anterior margin of the pectoral-fin spine mostly serrated). - -Parauchenoglanis patersoni - -is distinguished from - -P. stiassnoae - -by external mandibular barbel not reaching the tip of pectoral-fin spine (vs.reaching beyond the tip of pectoral-fin spine) and eyes positioned dorsolaterally (vs. dorsal). - -Parauchenoglanis patersoni - -is distinguished from - -P. ngamensis - -, - -P. lueleensis - -, - -P. poikilos - -, - -P. ernstswartzi - -, and - -P -. -megalasma - -by absence of black background spots in between the vertical bars or rows of spots [ -Fig. 11A -; vs. background spots present in - -P. ngamensis - -( -Fig. 5A -), - -P. lueleensis - -( -Fig. 10A -), and - -P. poikilos - -( -Fig. 11A -) or vermiculated pattern in - -P. ernstswartzi - -( -Fig. 12A -), or blotches in - -P -. -megalasma - -( -Fig. 13B -)]. It is further distinguished from these five species by absence or presence of only few spots on the head and fins (vs. heavily spotted in the other species). Furthermore, - -Parauchenoglanis patersoni - -is distinguished from - -P. dolichorhinus - -by a moderately depressed head, head depth 48.5%–54.9% HL (vs. depressed head, head depth 34.6%–40.0% HL); a deep body, 15.5%–21.4% SL (vs. shallower, 11.6%–14.0% SL), and posterior edge of anal fin reaching the posterior base of adipose fin (vs. not reaching posterior base of adipose fin). - -Parauchenoglanis patersoni - -is distinguished from - -P. chiumbeensis - -by posterior edge of anal fin reaching the posterior base of adipose fin (vs. reaching beyond level of posterior base of adipose fin) and external mandibular barbel not reaching tip of pectoral-fin spine (vs. reaching tip of pectoral-fin spine). - -Parauchenoglanis patersoni - -is distinguished from - -P. luendaensis - -by external mandibular barbel not reaching the tip of pectoral-fin spine (vs. reaching distal tip of pectoral-fin spine) and rounded caudal fin (vs. truncate). - - - - -Description: -Morphometric and meristic data are given in -Table 3 -. Body elongated. Dorsal body profile gently rising from snout tip to origin of dorsal fin, slightly concave from dorsal to adipose fin, and straight from adipose-fin origin to caudal-fin base. Body depth highest at origin of dorsal fin. Ventral body profile slightly convex from pectoral-fin origin to pelvic-fin origin and straight from pelvic-fin origin to caudal-fin base. Caudal peduncle laterally compressed. Anus and urogenital opening positioned about halfway between pelvic- and anal-fin origins. Adipose fin longer than anal-fin base, originating anteriorly to anal-fin origin, between pelvic-fin base and anal-fin origin, ending posteriorly to end of anal-fin base. Dorsal and pectoral fins with strong spine. Dorsal-fin origin anterior to pelvic-fin origin. Posterior edge of pectoral-fin ray does not reach pelvic fin. Anterior margin serrated on the distal end, and entire posterior margin of the pectoral-fin spine serrated. Caudal fin rounded. - - -Head moderately depressed. Snout profile bluntly triangular, on dorsal view ( -Fig. 11B -). Mouth subterminal. Lips fleshy. Eyes small and situated dorsolaterally. Three pairs of barbels, base thick and tips pointed. External mandibular barbel longest, almost reaching the tip of the pectoral-fin spine. Inner mandibular barbel shortest, reaching middle of eye. Maxillary barbel reaching posterior edge of the eye, sometimes reaching slightly beyond posterior edge of the eye. Posterior nostril slits positioned about halfway between snout tip and eye. - - -Colouration in life: -Overall body colouration varies from yellow dorsally to orange laterally. Mouth and distal tip of barbels and fins reddish (Supporting Information, -Fig. S2 -). Arrangement of spots is discussed below. - - -Colouration in alcohol: -Body light brown to grey dorsally and laterally, and light brownish to white ventrally. Body with five to six bars in small- and medium-sized specimens ( -42.9–62.6 mm -SL). Large-sized specimens (116.1–223.0 mm SL) with black spots smaller than the eye forming five to six (seven in -one specimen -) vertical rows. Additional black spots between bars or vertical rows generally absent, but a few scattered inconspicuous black spots might be present between the bars or vertical rows in both medium- and large-sized specimens. Head light brown. Fins with a few black spots, sometimes unspotted. Distal tip of dorsal fin rays with a narrow black margin. Usually, one or two small black spots present above gill opening and one spot on midline of caudal-fin base. Barbels unspotted. - - - - - -Distribution: -Parauchenoglanis patersoni - -is currently known from the mainstream of the Kwanza Basin ( -Angola -) above and below the Quissaquina Falls ( -Fig. 1 -). This species co-occurs in this river section with the other Kwanza endemic species, - -P. dolichorhinus - -. - - - - -Etomologo: -The species is named after Dr -Angus -Paterson, the former managing director of the National Research Foundation– South African Institute for Aquatic Biodiversity (NRF-SAIAB), in appreciation and recognition of his determination and efforts to build taxonomic expertise and drive ichthyological exploration in poorly surveyed areas in southern Africa. - - - - \ No newline at end of file diff --git a/data/03/C3/87/03C387D2FFD2365452849A58FC4DF9D1.xml b/data/03/C3/87/03C387D2FFD2365452849A58FC4DF9D1.xml new file mode 100644 index 00000000000..a8690363b92 --- /dev/null +++ b/data/03/C3/87/03C387D2FFD2365452849A58FC4DF9D1.xml @@ -0,0 +1,246 @@ + + + +A new booid snake from the Eocene (Lutetian) Konservat-Lagerstätte of Geiseltal, Germany, and a new phylogenetic analysis of Booidea + + + +Author + +Palci, Alessandro +School of Biological Sciences, University of Adelaide, Adelaide, SA 5000, Australia & South Australian Museum, North Terrace, Adelaide, SA 5000, Australia +alessandro.palci@adelaide.edu.au + + + +Author + +Onary, Silvio +Laboratório de Paleontologia, Faculdade de Filosofia Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Avenida Bandeirantes 3900, 14040 - 901, Ribeirão Preto, São Paulo, Brazil + + + +Author + +Lee, Michael S. Y. + + + +Author + +Smith, Krister T. +Department of Messel Research and Mammalogy, Senckenberg Research Institute, Senckenberganlage 25, 60325 Frankfurt am Main, Germany & Faculty of Biological Sciences, Goethe University, Max-von-Laue-Str. 9, 60438 Frankfurt, Germany + + + +Author + +Wings, Oliver +Natural History Museum, Fleischstr. 2, 96047 Bamberg, Germany & Natural Sciences Collections, Martin Luther University Halle-Wittenberg, Domplatz 4, 06108 Halle (Saale), Germany + + + +Author + +Rabi, Márton +Natural Sciences Collections, Martin Luther University Halle-Wittenberg, Domplatz 4, 06108 Halle (Saale), Germany & Department of Geosciences, Eberhard Karls Universität Tübingen, Hölderlinstr. 12, Tübingen 72074, Germany + + + +Author + +Georgalis, Georgios L. +Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Sławkowska 17, 31 - 016 Kraków, Poland + +text + + +Zoological Journal of the Linnean Society + + +2024 + +2023-12-12 + + +202 + + +2 + + +1 +27 + + + + +https://doi.org/10.1093/zoolinnean/zlad179 + +journal article +10.1093/zoolinnean/zlad179 +0024-4082 +14542189 +5DD1A6B-57F4-4DFF-9883-E7A684294FFFCorresponding + + + + + + +Genus + +Eoconstrictor +Scanferla & Smith + +, 2 0 2 0a + + + + + + + +Type +species: + + +Palaeopython fischeri +Schaal, 2004 + +(original combination), currently + +Eoconstrictor fischeri +( +Schaal, 2004 +) + +following the work of +Scanferla and Smith (2020a) +. + + + + + +Type +species locality and age: + +Messel Pit, Germany ( + +Smith +et al +. 2018 + +). All known specimens of + +E. fischeri + +come from the lacustrine ‘oil-shale’ of the Middle Messel Formation (near the Ypresian–Lutetian boundary, ~48 Mya) ( + +Lenz +et al +. 2015 + +). + + + + + +Emended diagnosis: +Eoconstrictor + +can be distinguished from all other snakes in having the following combination of features: edentulous premaxilla with bifid vomerine processes (unique autapomorphy); prootic with dorsoventrally compressed opening for exit of maxillary branch of trigeminal nerve ( +V +2), posterior margin of this foramen pointed (unique autapomorphy); between one and four labial foramina on the maxilla; 15–18 maxillary teeth; palatine with 5–6 teeth; pterygoid with 10–11 teeth; dentary with 17–19 teeth; mid-sagittal keel along the ventral side of the basioccipital contributing to V-shaped cross-section (adult feature, may be absent in juveniles); ectopterygoid process of pterygoid merging anteriorly with dentigerous process via gently concave (almost straight) margin (i.e. no deep emargination between dentigerous ramus and ectopterygoid process). This feature, in combination with a broad basipterygoid flange located immediately opposite on the medial side, gives the mid-portion of the pterygoid a broad, diamond-shaped to sub-elliptical appearance in dorsoventral view; total vertebral count up to 369 vertebrae, of which +c +. 246–303 are precloacal vertebrae, three are cloacal vertebrae (three vertebrae with lymphapophyses are visible in + +E. fischeri +SMF-ME 1607 + +), and up to +c +. 71 are caudal vertebrae. + +Eoconstrictor + +can be differentiated from + +Palaeopython + +Rochebrune, 1880 +, in having a thinner zygosphene bearing a prominent median lamellar tubercle, and in lacking a palatine foramen and a sigmoidal lateral margin of the maxilla (in dorsal view). + +Eoconstrictor + +can be differentiated from + +Paleryx +Owen, 1850 + +, in lacking a palatine foramen, in having a comparatively taller neural spine (especially on posterior trunk vertebrae), and in lacking a depressed neural arch on posterior trunk vertebrae. + +Eoconstrictor + +further differs from + +Phosphoroboa +Georgalis + +, Rabi & +Smith, 2021 +, in having a lower pterygoid tooth count (10–11 vs. 14), in lacking an anteromedial projection of the pterygoid at the palatine articulation, and in having a generally U-shaped frontoparietal suture (rather than V-shaped). + + + + +Remarks: +The number of precloacal vertebrae is best known in the type species of the genus, + +E. fischeri + +(cf. +Schaal 2004 +and +Smith and Scanferla 2022 +). +SMF-ME +1004 and +SMF-ME +2504 have a very high number of vertebrae (around 300 precloacals in +SMF-ME +2504), whereas +SMF-ME +1607 has notably fewer (around 246 precloacals). Given the stratigraphic distribution of these specimens, the variation does not seem to indicate secular change through time. +SMF-ME +1002 is from +1–3 m +above marked bed Alpha, +SMF-ME +2504 is from 6– +3 m +below Alpha (at +c +. 7000 yr/m, +28,000 +–63,000 +years apart), and +SMF-ME +1607 is from in between. + + +Based on data from +Szyndlar and Georgalis (2023) +on the minimum and maximum number of precloacal vertebrae in non-Caenophidian snakes (Supporting Information, File S3), precloacal variability [calculated as (Max—Min)/Min, where Max = maximum number of precloacals and Min = minimum number of precloacals] ranges between 0 and 0.5. + +Eoconstrictor fischeri + +, with a variability of (303 – 246)/246 = 0.23, would thus be close to the median value (Supporting Information, File S1: +Fig. S3 +). + + + + \ No newline at end of file diff --git a/data/03/C3/87/03C387D2FFD23658518D996EFE4CFAA1.xml b/data/03/C3/87/03C387D2FFD23658518D996EFE4CFAA1.xml new file mode 100644 index 00000000000..5d69ea44ee6 --- /dev/null +++ b/data/03/C3/87/03C387D2FFD23658518D996EFE4CFAA1.xml @@ -0,0 +1,787 @@ + + + +A new booid snake from the Eocene (Lutetian) Konservat-Lagerstätte of Geiseltal, Germany, and a new phylogenetic analysis of Booidea + + + +Author + +Palci, Alessandro +School of Biological Sciences, University of Adelaide, Adelaide, SA 5000, Australia & South Australian Museum, North Terrace, Adelaide, SA 5000, Australia +alessandro.palci@adelaide.edu.au + + + +Author + +Onary, Silvio +Laboratório de Paleontologia, Faculdade de Filosofia Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Avenida Bandeirantes 3900, 14040 - 901, Ribeirão Preto, São Paulo, Brazil + + + +Author + +Lee, Michael S. Y. + + + +Author + +Smith, Krister T. +Department of Messel Research and Mammalogy, Senckenberg Research Institute, Senckenberganlage 25, 60325 Frankfurt am Main, Germany & Faculty of Biological Sciences, Goethe University, Max-von-Laue-Str. 9, 60438 Frankfurt, Germany + + + +Author + +Wings, Oliver +Natural History Museum, Fleischstr. 2, 96047 Bamberg, Germany & Natural Sciences Collections, Martin Luther University Halle-Wittenberg, Domplatz 4, 06108 Halle (Saale), Germany + + + +Author + +Rabi, Márton +Natural Sciences Collections, Martin Luther University Halle-Wittenberg, Domplatz 4, 06108 Halle (Saale), Germany & Department of Geosciences, Eberhard Karls Universität Tübingen, Hölderlinstr. 12, Tübingen 72074, Germany + + + +Author + +Georgalis, Georgios L. +Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Sławkowska 17, 31 - 016 Kraków, Poland + +text + + +Zoological Journal of the Linnean Society + + +2024 + +2023-12-12 + + +202 + + +2 + + +1 +27 + + + + +https://doi.org/10.1093/zoolinnean/zlad179 + +journal article +10.1093/zoolinnean/zlad179 +0024-4082 +14542189 +5DD1A6B-57F4-4DFF-9883-E7A684294FFFCorresponding + + + + + + + +Eoconstrictor barnesi + +sp.nov. + + + + + + + +( +Figs 2–1 +0, Supporting Information, File S1: +Figs S1 +, S +2 +) + + + +Zoobank registration: +This published work and the nomenclatural act it contains have been registered in ZooBank, the official registry of zoological nomenclature for the International Code of Zoological Nomenclature (ICZN). The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix ‘https://zoobank.org/’. The LSID for this publication is: + +urn:lsid:zoobank.org:pub: +7EA 0164B-217B-49AD-88FC-20F231060251 + +, while the LSID for the new species + +Eoconstrictor barnesi + +is: + +urn:lsid:zoobank.org: act: +75DD1A6B-57F4-4DFF-9883-E7A684294FFF + + + + + +Figure 2. + +Eoconstrictor barnesi +, GMH + +XXXVIII-20-1964, holotype. A, View of the specimen as preserved, insets show clusters of vertebrae that are preserved on the same slab but a few centimetres away from the rest of the body; (B) close-up of the skull in dorsal view and disarticulated left lower jaw in lateral view; (C) close-up of three mid-trunk vertebrae in ventral view; (D) close-up of two posterior trunk vertebrae in ventral view; (E) close-up of four mid-trunk vertebrae in left lateral view; (F) close-up of four mid-trunk vertebrae in dorsal view. + + + + +Figure 3. + +Eoconstrictor barnesi +, GMH + +XXXVIII-20-1964, holotype, digital renderings of the segmented skull. A, skull in dorsal view and disarticulated left lower jaw in lateral view. B, skull in ventral view and disarticulated left lower jaw in medial view. Abbreviations: c, coronoid; co, compound; d, dentary; f, frontal; j, fragment of jugal?; ma, maxilla; p, parietal; pa, palatine; pt, pterygoid; sp, splenial; st, supratemporal. + + + + +Etymology: +Species epithet honouring +Ben +Barnes, who pioneered palaeontological excavations and research in the brown coal deposits of Geiseltal in the 1920s, as part of his Ph.D. studies at the Geological Institute of Halle (Saale). + + + + + + +Holotype +: + +GMH XXXVIII-20-1964 +, a partial skull and partially disarticulated skeleton missing only the most anterior precloacal region, the cloacal region, and most of the tail ( +Figs 2–5 +, Supporting Information, File S1: +Fig. S1 +). + + + + + +Paratype +: + +GMH LIX-3-1992 +, a moderately complete and only slightly disarticulated skull and skeleton preserving the anterior half of the body ( +Figs 6–10 +, Supporting Information, File S1: +Fig. S2 +) + +. + + +Type locality and age: +The +Konservat-Lagerstätte of Geiseltal +, located about + +30 km +west of Leipzig + +and + +20 km +south-east of Halle + +, in the state of +Saxony-Anhalt +, +Germany +( +Barnes 1927 +, + +Krumbiegel +et al +. 1983 + +). The fossils were recovered from fossil sites within coal mines identified by Roman numerals (e.g. LIX, XXXVIII). Both fossil specimens come from fossil sites in the upper middle coal seam (obere Mittelkohle). The age of the deposits is broadly estimated as +late early or middle Eocene +(~Lutetian, i.e. anywhere between 41 and 48 Mya) ( + +Georgalis +et al. +2021 + +). + + + + + +Diagnosis: +Eoconstrictor barnesi + +sp. nov. +can be distinguished from + +E. fischeri + +based on the presence of a single labial foramen in the maxilla (four in + +E. fischeri + +), the different position of the mental foramen, which is located ventral to the 4 +th +tooth position (ventral to the 5 +th +tooth position in + +E. fischeri + +), and based on the lateral offset of the mandibular cotyle relative to the rest of the compound bone (offset absent in + +E. fischeri + +). + +Eoconstrictor barnesi + +shares the presence of a single labial foramen in the maxilla with + +E. spinifer + +, but can be distinguished from the latter based on: its more elongated maxilla (length/height ratio between 7.5 and 7.9 vs. a ratio of ~ +4.68 in + +E. spinifer + +; the maxilla of + +E. spinifer + +is slightly damaged posteriorly, but based on its posterior tapering no more than two tooth positions are missing); a quadrate ramus of the pterygoid that is gently curved laterally (forming a distinct ~120° angle in + +E. spinifer + +); the presence of a median tubercle on the zygosphene (absent in + +E. spinifer + +, at least in the anterior precloacal vertebrae, more posterior vertebrae are unknown in that species); an almost straight anterior margin of the zygosphene (excluding lateral projections) [crenate +sensu +Auffenberg (1963) +in + +E. spinifer + +]; and presence of prezygapophyseal accessory processes throughout the precloacal vertebral column (absent at least in the anterior precloacal vertebrae in + +E. spinifer + +). + +Eoconstrictor barnesi + +differs from both + +E. fischeri + +and + +E. spinifer + +in the presence of small paracotylar foramina on at least some of its vertebrae and in the anterior margin of the foramen for the exit of the mandibular branch of the trigeminal (V3), which is gently rounded instead of sharply pointed. Although + +E. fischeri + +and + +E. spinifer + +share a ventromedian crest on the basioccipital ( +Scanferla and Smith 2020a +, + +Georgalis +et al +. 2021 + +), this element is currently unknown in + +E. barnesi + +. + + + + +Description + + +Cranial material of + +GMH +XXXVIII-20-1964 +( +holotype +) + + + +Based on the relative length of the lower jaw, the skull of this specimen is about 35% larger than that of the paratype ( +GMH +LIX-3-1992 +) but unfortunately far less complete ( +Figs 2B +, +3 +). The only bones that are present and not too badly crushed are the frontals, parietal, left jugal, right supratemporal, right maxilla, left palatine and pterygoid, most of the left lower jaw (missing only the angular), and the right dentary. + + +The frontals are fairly well preserved, except ventrally, where the descending flanges and medial pillars are broken off ( +Fig. 4A–D +). However, a portion of the medial frontal pillars is still preserved dorsally, indicating that these were present. In dorsal view each frontal has an almost semicircular outline, with a distinct lateral flange above the orbit. In lateral view, the frontals show distinct articular surfaces for the prefrontals and the parietal, and the descending flanges, although partially damaged, were clearly deeper posteriorly. + + + +Figure 4. + +Eoconstrictor barnesi +, GMH + +XXXVIII-20-1964, holotype, digital renderings of the segmented skull bones. A–D, frontals in dorsal (A), ventral (B), anterior (C), and left lateral (D) view; E–F, possible fragment of left jugal in lateral (E) and medial (F) view; G, right supratemporal in lateral view; H–K, parietal in dorsal (H), ventral (I), left lateral (J), and anterior (K) view; L–O, right maxilla in dorsal (L), lateral (M), medial (N), and ventral (O) view; P–S, left pterygoid in dorsal (P), ventral (Q), lateral (R), and medial (S) view; T–W, left palatine in dorsal (T), ventral (U), lateral (V), and medial (W) view. Abbreviations: asaf, anterior superior alveolar foramen; bv, blood vessel; chp, choanal process; ep, ectopterygoid process of pterygoid; flf, frontal lateral flange; fmp, frontals median pillars; malf, maxillary labial foramen; mp, maxillary process; mgqr, medial groove on quadrate ramus; mV2, maxillary foramen for maxillary branch of trigeminal nerve (V2); mV2a, accessory foramen for maxillary branch of trigeminal nerve (V2); pmc, parietal midsagittal crest; pmmc, posteromedial maxillary crest; pp, palatine process of maxilla; mV2, maxillary foramen for maxillary branch of trigeminal nerve (V2); qr, quadrate ramus of pterygoid. + + + + +Figure 5. + +Eoconstrictor barnesi +, GMH + +XXXVIII-20-1964, holotype, digital renderings of the segmented lower jaw bones. A–C, left dentary in dorsal (A), lateral (B), and medial (C) view; (D) left dentary in medial view with associated splenial in articulation; (E–G) left compound bone in dorsal (E), lateral (F), and medial (G) view; (H) splenial in medial view; (I–J) left coronoid in medial (I) and lateral (J) view. Abbreviations: aesc, crest for attachment of adductor externus superficialis muscle on compound bone; af, adductor fossa; asf, anterior surangular foramen; cp, coronoid process; pdp, posterodorsal dentary process; mc, mandibular condyle; mf, mental foramen; mg, Meckelian groove; sac, surangular crest; sap, surangular process; sp, splenial. + + + +Unlike the paratype, this specimen also preserves the ventral portion of the left jugal ( +sensu +Palci and Caldwell 2013 +). The dorsal part that articulated between the frontal and the parietal is missing ( + +Figs +3 + +, +4E–F +). The preserved part of the element has a claw-shaped profile, with a broad dorsal end (incomplete) and a tapering descending ramus that is recurved posteriorly. The element closely resembles the ventral half of the jugal in + +E. fischeri +SMF-ME + +11 398. + + +The right supratemporal is very similar to that of the paratype, has the same dorsomedial curvature, expanded and gently rounded anterior end, and thick posterior end bearing a broad facet for articulation with the quadrate ( +Fig. 4G +). + + +The parietal is nearly complete and very robust ( +Fig. 4H–K +). Compared to the parietal of the paratype it is relatively broader anteriorly, and more triangular in dorsal view. A distinct mid-sagittal crest is present on the posterior half of this bone and is replaced anteriorly by a round crest that merges into a subtriangular mound just posterior to the frontoparietal suture. The frontoparietal suture appears straighter in the holotype compared to the paratype, where it is clearly U-shaped (i.e. distinctly concave anteriorly). This difference is unlikely to be an artefact of poor preservation of the anterior margin of the parietal, since the only gently curved posterior margin of the frontals matches the anterior margin of the parietal, indicating that this is the genuine morphology present in the holotype. + + +The right maxilla is quite robust but still relatively elongate (length to height ratio = 7.5). Its ventral margin is straight and bears 17 tooth positions, whereas the dorsal margin has a distinctive sigmoidal curvature in lateral view ( +Fig. 4L–O +), which is absentintheparatypeGMHLIX-3-1992(notethatthissigmoidal curvature is not the same as that observed in Messelopythonidae, where the curvature affects the lateral margin of the maxilla and is observed in dorsal view; +Smith and Scanferla 2022 +). However, as in the paratype, the maxilla of GMH XXXVIII-20-1964 also bears an anterior superior alveolar foramen that is located dorsal to the third maxillary tooth position, and has two foramina for the entry of the maxillary division of the trigeminal nerve (V2). The main, larger entrance of the trigeminal nerve in the maxilla is located on the medial end of the palatine process, whereas a foramen for a secondary smaller branch is located just posterior to this process, near its base. The palatine process is quadrangular in shape and anteromedially long. As in the paratype, the maxilla of GMH XXXVIII-20-1964 also has a thin medial crest running along the posteromedial margin, where this bone would articulate with the ectopterygoid. In dorsal view the maxilla is relatively straight posteriorly, and gently curved medially in the anterior region. + + +The left pterygoid is relatively well preserved and only lacks the distal half of the quadrate ramus and a small fragment just posterior to the dentigerous portion ( +Fig. 4P–S +). The dentigerous ramus preserves 10 tooth positions and has a concave medial margin. An additional tooth may have been present at the end of the dentigerous ramus, where the bone is damaged.The pterygoid is broadest at the level of the ectopterygoid process, which is quite robust and terminates in a broad sub-elliptical facet for articulation with the ectopterygoid. The longitudinal axis of this facet is tilted anterodorsally. On the opposite side of the ectopterygoid process, the pterygoid expands into a broad rounded flange (basipterygoid flange). The quadrate ramus is relatively deep, at least anteriorly where it is preserved, and bears a deep medial longitudinal groove for the m. protractor pterygoidei. + + + +Figure 6. + +Eoconstrictor barnesi +, GMH + +LIX-3-1992, paratype. A, View of the specimen as preserved; (B) close-up of the skull in left lateral view; (C) close-up of three precloacal vertebrae in ventral view; (D) close-up of five precloacal vertebrae in left dorsolateral and dorsal views. + + + + +Figure 7. + +Eoconstrictor barnesi +, GMH + +LIX-3-1992, paratype, digital renderings of the segmented skull. A, skull in left lateral view; (B) skull in dorsal view; (C) skull in right ventrolateral view. Abbreviations: a, angular; c, coronoid; co, compound; d, dentary; ec, ectopterygoid; f, frontal; ma, maxilla; n, nasal; p, parietal; pa, palatine; pf, prefrontal; pr, prootic; pt, pterygoid; q, quadrate; sp, splenial; st, supratemporal. + + + + +Figure 8. + +Eoconstrictor barnesi +, GMH + +LIX-3-1992, paratype, digital renderings of the segmented skull bones. A–C, nasals in dorsal (A), lateral (B), and anterior (C) view; (D–F) left prefrontal in dorsal (D), lateral (E), and posterior (F) view; (G–K) frontals in dorsal (G), ventral (H), anterior (I), posterior (J), and left lateral (K) view; (L–O) parietal in dorsal (L), ventral (M), anterior (N), and left lateral (O) view; (P–Q) supratemporals in dorsal (P) and left lateral (Q) view; (R) prootic in left lateral view; (S–T) right quadrate in lateral (S) and posterior (T) view; (U–X) left maxilla in dorsal (U), lateral (V), medial (W), and ventral (X) view; (Y–Z, A’) left palatine in dorsal (Y), lateral (Z), and ventral (A’) view; (B’) ectopterygoid in dorsal view; (C’) overlapping pterygoids, right pterygoid in ventrolateral view obscuring most of left pterygoid lying underneath (in dorsolateral view). Abbreviations: alp, anterolateral process of ectopterygoid; amp, anteromedial process of ectopterygoid; asaf, anterior superior alveolar foramen of maxilla; bv, foramen for blood vessel; chp, choanal process of palatine; cid, foramina for exit (posterior) and re-entry (anterior) of nerve that innervates the constrictor internus dorsalis musculature; df, descending flange of nasal; dl, dorsal lappet of prefrontal; ep, ectopterygoid process of pterygoid; fmp, frontals median pillars; fpf, facet for articulation with prefrontal; fp, facet for articulation with parietal; fq, facet for articulation with quadrate; fst, facet for articulation with supratemporal; hl, horizontal lamina of nasal; lf, lacrimal foramen; ll, lateral lamina of prefrontal; malf, maxillary labial foramen; mp, maxillary process of palatine; mV2a, maxillary foramen for accessory branch of maxillary division of trigeminal nerve (V2); mV2, maxillary foramen for maxillary division of trigeminal nerve (V2); pmc, parietal midsagittal crest; pmmc, posteromedial maxillary crest; pp, palatine process of maxilla; qr, quadrate ramus of pterygoid; V2, exit for maxillary branch of trigeminal nerve (cranial nerve V); V3, exit for mandibular branch of trigeminal nerve; VIIh, exit for hyomandibular branch of facial nerve (cranial nerve VII); VIIp, exit for palatine branch of facial nerve. + + + + +Figure 9. + +Eoconstrictor barnesi +, GMH + +LIX-3-1992, paratype, digital renderings of the segmented lower jaw bones. A–C, left dentary in dorsal (A), lateral (B), and medial (C) view; (D–F) left compound bone in dorsal (D), lateral (E), and medial (F) view; (G) right splenial and angular in medial view; (H) right splenial and angular in lateral view (i.e. showing sutural contacts with dentary and compound, respectively); (I–J) left coronoid in medial (I) and lateral (J) view. Abbreviations: a, angular; aesc, crest for attachment of adductor externus superficialis muscle on compound bone; af, adductor fossa; aif, anterior intermandibularis nerve foramen; asf, anterior surangular foramen; cp, coronoid process; mc, mandibular condyle; mf, mental foramen; mg, Meckelian groove; pac, prearticular crest; pdp, posterodorsal process of dentary; pif, posterior intermandibularis nerve foramen; sac, surangular crest; sap, surangular process; sp, splenial. + + + +The left palatine bears six tooth positions and a distinct maxillary process posterolaterally ( +Fig.4T–W +). This process is short and square in shape, and slightly damaged distally. The choanal process lies on the opposite side, and appears to be very short. There is the possibility that the process may be broken off distally, but its similarity to the same process in the paratype suggests that this may in fact be the actual morphology of the bone. A distinct ridge runs dorsally along the anteromedial edge of the choanal process and merges anteriorly with the dorsal surface of the dentigerous process of the palatine.A distinct concave facet for a tongue-in-groove articulation with the pterygoid is visible at the posterior end of the palatine in ventral view. In medial and lateral views, the palatine is deeper at mid-length and its dorsal margin slopes gently anteriorly and posteriorly. There is no foramen for the passage of a branch of the trigeminal nerve. This palatine is overall very similar to an isolated palatine (YPM-VPPU 12281) from the Phosphorites du Quercy, +France +, figured + +in Georgalis +et al. +(2021 + +: fig. 105). + + + +Lower jaw of +GMH +XXXVIII-20-1964 +( +holotype +) + + + +The lower jaw of this specimen is reasonably complete, missing only the angular ( +Fig. 5 +). The left dentary is the better preserved of the two elements ( +Figs 3B +, +5A–C +). It is fairly robust, slightly arched in lateral view and mostly straight in dorsal view, except towards its anterior end, where it bends medially to a small degree. A relatively small mental foramen is located ventral to the fourth tooth position. There is a distinct posterodorsal ramus, deeper and twice the length of the posteroventral ramus. The two rami are separated by a deep notch where the surangular process of the compound bone articulates. The Meckelian groove is shallow, runs along the ventral margin of the dentary and extends anteriorly to its far end. The dentary bears 18 tooth positions. + + + +Figure 10. + +Eoconstrictor barnesi +, GMH + +LIX-3-1992, paratype, digital renderings of the segmented precloacal vertebrae. A–E, anterior precloacal in anterior (A), posterior (B), left lateral (C), dorsal (D), and ventral (E) view; (F–J) middle precloacal in anterior (F), posterior (G), left lateral (H), dorsal (I), and ventral (J) view. + + + +The compound bone ( +Fig. 5E–G +) is slightly longer than the dentary and bears a prominent rounded coronoid process with an anterior margin that forms almost a 90° angle with the straight dorsal margin of the surangular process. At the base of the coronoid process, where this meets the surangular process, lies the anterior surangular foramen. A tall surangular crest slopes posteroventrally from the coronoid process and delimits the adductor fossa laterally. The prearticular crest is somewhat damaged, but appears to be much lower than the surangular crest. Posteriorly, the mandibular cotyle is delimited anteriorly by a distinct transverse crest, which laterally merges with the crest for the insertion of the m. adductor externus superficialis. This latter crest slopes anteroventrally from its position near the cotyle until it approaches the ventral margin of the compound bone halfway toward the coronoid process, then gently rises again until it reaches a point ventral to the coronoid process. + + +The left splenial is preserved and appears to be missing only its most posterodorsal portion ( +Fig. 5D, H +). It is an elongate, subtriangular element, with a distinct longitudinal medial keel that fits into the Meckelian groove. A foramen for the anterior intermandibularis nerve is only partially preserved, the portion of the bone framing the foramen dorsally being lost due to breakage. The articular surface for the angular is only partially preserved (dorsal portion missing). + + +The left coronoid is complete, and has a distinct hockeystick shape, with a strap-like ascending process expanding anteroventrally into a broader, mediolaterally flat anteroventral portion ( +Fig. 5I, J +). The angle between the posterior ascending ramus and the horizontal portion is about 120°. + + + +Postcranial material of GMH XXXVIII-20-1964 ( +holotype +) + +The postcranial skeleton of this specimen is partially disarticulated into a series of segments, but a large part of the body is preserved, with the exception of the most anterior precloacal, the cloacal, and most of the caudal regions. Only about 10 distal caudal vertebrae remain ( +Fig. 2 +; Supporting Information, File S1: +Fig. S1 +). This snake has at least 189 vertebrae preserved, but if we consider that the most anterior elements, the cloacal region, and most of the tail are missing (and possibly also the posterior end of the precloacal series), then the total number of vertebrae must have been much higher. In fact, the number of precloacal vertebrae in specimens of + +E. fischeri + +varies between 246 and 303 (see Remarks above), so it is plausible that + +E. barnesi + +had a similar number, with some variability among individuals. + + +The average centrum length of the largest vertebrae in GMH XXXVIII-20-1964 is +12.8 mm +, and the vaulting ratio ( +sensu + +Georgalis +et al +. 2021 + +) is 0.33 (averages from 4 among the largest mid-trunk vertebrae). + + +The overall vertebral morphology of this specimen is mostly consistent with that observed in members of +Boidae +(see +Szyndlar and Georgalis 2023 +). The anterior trunk vertebrae have distinct hypapophyses that decrease in height posteriorly and eventually turn into low but distinct haemal keels on the mid-trunk vertebrae. Posteriorly in the trunk region, the haemal keel grades into a gently rounded surface with a small posteroventral tubercle. The most posterior precloacal vertebrae have a very narrow, mediolaterally compressed, mid-sagittal portion of the centrum that is delimited laterally by deep subcentral fossae. There are only 10 posterior caudal vertebrae that can be confidently identified, four preserved in dorsal view and six in left lateral view ( +Fig. 2 +; Supporting Information, File S1: +Fig. S1 +). These vertebrae are anteroposteriorly elongate, bear low neural spines, small but distinct prezygapophyseal accessory processes, and well-developed subtriangular haemapophyses that point posteroventrally. + + +All vertebrae possess a pair of small subcentral foramina, at least in all those where this feature can be ascertained. The neural spines of the mid-trunk vertebrae are trapezoidal in lateral view, dorsally flat and with an anterior edge that is well posterior to the zygosphene roof. Posteriorly they slightly overhang the condyle, which is robust and hemispherical and otherwise exposed in dorsal view in disarticulated vertebrae. Prezygapophyseal accessory processes can be observed in all of the preserved regions of the body. Parasagittal ridges +sensu + +Hsiou +et al +. (2014) + +are present on the most robust mid-trunk vertebrae, and at their posterior end sometimes is a small, rounded tubercle, likely marking the insertion of a tendon from the transversospinalis muscle complex. + +The zygosphene on mid-trunk vertebrae is relatively thin (width to height ratio = 3.2, where width is measured between the dorsolateral corners of the zygosphene and height is measured in the middle of the zygosphene) and bears a median tubercle. The median tubercle is subtriangular in dorsal view and dorsoventrally thin (lamellar). The anterior margin of the zygosphene is almost straight (excluding lateral projections). + +Based on microCT scan images, paracotylar foramina appear to be present on at least some of the vertebrae, but due to the small size of the foramina and the numerous fractures affecting most of the vertebrae it is unclear how frequent this feature is across the vertebral column. Note that paracotylar foramina are present in some (but not all) members of +Boidae +(see +Szyndlar and Georgalis 2023 +). + +The ribs are distinctly curved, with a posterodorsal tubercle and a small foramen on the shaft distal to the articular head. + + +Cranial material of +GMH +LIX-3-1992 +( +paratype +) + + + +The skull of this specimen is well preserved albeit not complete ( +Fig. 6B +). Most bones could be segmented from the microCT scan data and visualized digitally in three dimensions ( +Figs 7 +, +8 +). The premaxilla, basioccipital, supraoccipital, otoccipitals, parabasisphenoid, right prootic, vomers, septomaxillae, right maxilla, and right palatine are absent or too fragmentary to be digitally isolated. + + +Both nasals are partially preserved in this specimen ( +Fig. 8A–C +). The dorsal horizontal lamina is best preserved in the right nasal and has a triangular shape. The descending flange of the nasals is deeper anteroventrally. There is no indication of a process that may have contacted the frontals posteroventrally, and the contact must have been exclusively posterodorsal. + + +Both prefrontals are preserved ( +Fig. 7 +), but the left element is somewhat better preserved ( +Fig. 8D–F +). The prefrontal bears a broad subtriangular lateral lamina, which extends dorsally into a tongue-shaped dorsomedial lappet. The well-developed medial and lateral foot processes delimit a lacrimal foramen that is open ventrally. + + +Both frontals are present in this specimen, and preserved only slightly offset relative to one another ( +Fig. 8G–K +). In dorsal view they have a subrectangular to trapezoidal outline with a flat dorsal surface. Anteromedially they bear a protruding process for articulation with the nasals. Ventrally the frontals are in contact to fully enclose the space for the olfactory bulbs. Distinct frontal medial pillars are present and meet the lateral flanges to fully enclose the olfactory tracts ( +Fig. 8I +). In lateral view articular facets for the prefrontals and parietal are visible anteriorly and posterodorsally, respectively. The frontals are much deeper posteriorly, where they articulated with the missing parabasisphenoid. + + +The parietal is very well preserved ( +Fig. 8L–O +). Anteriorly it bears a U-shaped sutural margin for the frontals and has greatly expanded anterolateral processes for articulation with the jugals ( +sensu +Palci and Caldwell 2013 +), which unfortunately are not preserved in this specimen (note that there is some disagreement about the presence of jugals in modern snakes, with the elements in this position behind the orbit interpreted as postfrontals by + +Zaher +et al. +(2023) + +; however, see + +Segall +et al. +(2023) + +. In dorsal view the parietal tapers posteriorly forming an almost triangular outline. A distinct mid-sagittal crest is present on the dorsal surface and extends along the posterior half of the bone. In the anterior half the crest is replaced by a low ridge, which appears to extend anteriorly up to the fronto-parietal suture; however, this could be artifactual because the feature is not observed in the holotype (GMH XXXVIII-20-1964) and the anterior region of the parietal presents some fractures. + + +Both left and right supratemporals are preserved, with the former in slightly better condition ( +Fig. 8P, Q +). In dorsal view the supratemporals are slightly curved, with the concavity facing dorsomedially. The supratemporals are flattened and expanded anteriorly, where they articulate with the prootic and parietal, and taper and thicken posteriorly. The anterior margin of the better preserved supratemporal is gently rounded. A distinct broad facet for articulation with the dorsal condyle of the quadrate is present posterolaterally. + + +The left prootic is fairly well preserved ( +Fig. 8R +). This bone shows the two openings for the exit of the maxillary (V2) and mandibular (V3) divisions of the trigeminal nerve, separated by a laterosphenoid ossification (‘ophidiosphenoid’ +sensu +Gauthier +et al. +2012). The opening for cranial nerve V3 has a rounded anterior margin, unlike that of + +E. spinifer + +and + +E. fischeri + +, where the margin of the foramen is pointed anteriorly ( + +Georgalis +et al +. 2021 + +). The opening for the maxillary branch is not closed by the prootic anteriorly but by the parietal. A foramen for the exit of the hyomandibular branch of the facial nerve (cranial nerve VII) is visible just posterior to the exit of the mandibular branch of the trigeminal. Just ventral to this exit is a foramen for the palatine branch of the facial nerve. Foramina for the exit and re-entry of the nerve that innervates the constrictor internus dorsalis muscle complex (‘cid nerve’ of +Rieppel 1979 +) are visible along the anteroventral margin of the prootic. This margin slopes anterodorsally and forms the sutural contact with the missing parabasisphenoid. Posteroventrally the margin of the prootic slopes at a 45° angle and dorsally forms the anterior margin of the fenestra ovalis in a distinct crista prootica (part of the crista circumfenestralis). The dorsal margin of the prootic is subhorizontal and posterodorsally a shallow excavation marks the articular surface for the supratemporal. + + +Both left and right quadrates are preserved ( +Fig. 7 +), but the left quadrate is fractured at the middle of the shaft, and the two halves are slightly displaced relative to one another. The right quadrate is preserved in much better condition ( +Fig. 8S–T +). In lateral view, the right quadrate has a narrow, subtriangular outline, with a straight anterior margin and a short dorsal margin that slants posteroventrally. In posterior view the bone is constricted mediolaterally just above the ventral condyle, which is distinctly expanded and has a saddle-shaped articular surface for the compound bone. There is no clear indication of the presence of a stylohyal process, but this could be because of poor preservation and/or scan resolution. + + +The left maxilla is well preserved ( +Fig. 8U–X +). It is an elongate bone (length to height ratio = 7.9) bearing 16 tooth positions, but only seven teeth are preserved in position. There is no facial process, and the maxilla simply increases in depth anteriorly quite gently. In dorsal view an anterior superior alveolar foramen is clearly visible above the third tooth position. Three distinct foramina are present medial, anterior, and posterior to the palatine process. The anterior foramen (variably present in snakes) is here interpreted as the exit of a nerve and/or blood vessel, whereas the other two foramina are for the entry in the maxilla of branches of the maxillary division of the trigeminal nerve (cranial nerve V2). The palatine process is almost triangular in dorsal view, with a long anteromedial edge. Laterally the maxilla bears a single labial foramen located above the fourth tooth position. The ventral margin of the maxilla is straight. In dorsal and ventral views, the anterior end of the maxilla is gently bowed medially. There is no distinct ectopterygoid process, but a thin medial shelf extends along the posterior quarter of the bone. + + +Both palatines are preserved, but the right element is very badly crushed and could not be segmented. The left palatine, on the other hand, is nearly complete ( +Fig. 8Y, Z +, A’). The palatine lacks a foramen for the suborbital branch of the maxillary division of the trigeminal nerve and appears to have a short and thin choanal process. This could be due to poor preservation, and part of the process may be missing, but the same condition is also observed in the holotype. The anterior edge of the choanal process merges with the dorsal surface of the palatine along a thin crest. The maxillary process is complete, subrectangular in dorsal view, and slightly bent posterolaterally. There are six tooth positions. In lateral view the palatine is deeper at the level of the maxillary process, and its dorsal margin slopes down anteriorly and posteriorly from that point. + + +The ectopterygoid could be identified only on the left side ( +Fig. 8 +B’). It is a flat, V-shaped bone, with distinct medial and lateral processes for articulation with the maxilla. The medial process is much broader than the lateral one and convex in shape. The notch separating the two processes is likely artifactual. The posterior end of the bone is damaged and a surface for articulation with the pterygoid cannot be recognized. + + +Although both pterygoids are preserved, they lie on top of each other, which complicates the interpretation of their morphology ( +Fig. 8 +C’). The right pterygoid is the better preserved of the two. While an accurate tooth count is difficult to obtain on the right pterygoid due to the numerous fractures in this element, 11 tooth positions can be identified on the left pterygoid. The dentigerous portion is fairly straight, whereas the quadrate ramus arches posterolaterally, is blade-like in shape and has a distinct longitudinal groove for the attachment of the musculus (m.) protractor pterygoidei. A distinct and stout ectopterygoid process is present lateral to the posterior half of the tooth row; this process has a broad medial base and tapers into a laterally facing sub-elliptical articular facet for the ectopterygoid. + + + +Lower jaw of +GMH +LIX-3-1992 +( +paratype +) + + + +The left lower jaw of this specimen is very well preserved ( +Figs 6 +, +7 +). The left dentary ( +Fig. 9A–C +) is a robust element that bears 17 tooth positions, with 11 teeth still in place. In dorsal view this bone is broader posteriorly than anteriorly and is gently curved mediolaterally. A single mental foramen is present ventral to the fourth tooth position. The posterodorsal ramus of the dentary is deeper and about twice the length of the posteroventral ramus. In medial view, a shallow Meckelian groove extends all the way from the posterior notch between posterodorsal and posteroventral rami to the anterior end of the dentary. A medial subdental shelf is absent. + + +The compound bone ( +Fig. 9D–F +) is slightly longer than the dentary and bears a distinct surangular process that fits between the posterodorsal and posteroventral rami of the latter bone. The dorsal margin of the surangular process slopes posterodorsally into a distinct coronoid process. Anteroventral to this process, in lateral view, an anterior surangular foramen is visible. The adductor fossa is deep, well defined, and bordered medially and laterallybytheprearticularandsurangularcrests, respectively.These crests are subequal in height. The mandibular cotyle is slightly offset laterally relative to the rest of the compound bone, because of a gentle bend in the posterior quarter of this bone (this offset is absent in + +E. fischeri + +, where the cotyle is in line with the main body of the compound). A distinct mediolateral crest marks the anterior margin of the mandibular cotyle. There is effectively no retroarticular process, but a blunt ending of the compound posterior to the cotyle. The ventral margin of the compound bone is relatively straight, with only a gentle sigmoidal curvature. Along the ventrolateral margin of the compound bone is a longitudinal crest, presumably for the insertion of the m. adductor externus superficialis ( +Fig. 9E +). This crest starts anteriorly in the middle of the ventral margin of the compound and extends posteriorly up to the anterior margin of the mandibular cotyle. + + +The splenial and angular are not preserved on the left side, but fortunately their right counterparts are ( +Fig. 9G–H +). The right angular is complete, while the right splenial is missing the most anterior portion, but is preserved well enough that most details are visible. The intramandibular joint is clearly visible between the angular and the splenial, with a convex surface on the former fitting into a concave facet on the latter. Both elements are subtriangular, although the angular is much deeper at mid-length and slightly longer anteroposteriorly. A foramen for the exit of a branch of the intermandibularis nerve is visible on each element (i.e. anterior and posterior mylohyoid foramina). + + +The left coronoid is well preserved, although a small portion of its anteroventral end may be missing due to breakage ( +Fig. 9I, J +). This element is comma shaped and strap like and articulates with the medial side of the well-developed coronoid process on the compound. Both the coronoid bone and the coronoid process of the compound are subequal in height when articulated. + + + +Postcranial material of + +GMH +LIX-3-1992 +( +paratype +) + + + + +The postcranium of this specimen is preserved almost in articulation, with very little distortion and displacement ( +Fig. 6A +). Only the cloacal and caudal regions cannot be identified. There are at least 113 preserved precloacal vertebrae. The largest mid-trunk vertebrae are +10.8 mm +long (average centrum length of four among the largest mid-trunk vertebrae). + + +As in the holotype described above, the vertebral morphology of the paratype is consistent with that of +Boidae +(see +Szyndlar and Georgalis 2023 +). The anterior precloacals have long and slender hypapophyses, strongly projecting posteriorly, and equally long neural spines ( +Fig. 10 +). Their synapophyses project ventrally below the ventral margin of the small and subcircular cotyle. The mid-trunk vertebrae ( +Fig. 10F–J +) lack hypapophyses, but have a distinct mid-sagittal keel; the midventral keel disappears in the most posterior precloacals ( +Fig. 6C +). The ‘vaulting ratio’ ( +sensu + +Georgalis +et al +. 2021 + +) of the two segmented vertebrae is 0.55 for the anterior precloacal ( +Fig. 10A–E +) and 0.32 for the mid-trunk vertebra ( +Fig. 10F–J +). The centra are robust, subtriangular in shape, and bear a pair of small subcentral foramina, features that are similar to those of other booids. The condyles are robust and hemispherical, exposed in dorsal view in disarticulated vertebrae. The neural spines are anteroposteriorly short with the anterior edge well posterior to the zygosphene roof. In lateral view the neural spines are trapezoidal, with a slanting anterior margin and a straight dorsal margin. Parasagittal ridges ( +sensu + +Hsiou +et al +. 2014 + +) are present on the most robust mid-trunk vertebrae. Distinct small prezygapophyseal accessory processes are consistently present on all precloacal vertebrae. In dorsal view the anterior margin of the zygosphene is almost straight (excluding lateral projections) and is marked by the presence of a distinct sub-triangular median tubercle. The tubercle is broad mediolaterally at its base, but fairly thin dorsoventrally (lamellar tubercle +sensu + +Georgalis +et al +. 2021 + +). The zygosphene of the mid-trunk vertebrae is relatively thin (width to height ratio = 3.5). The two vertebrae that were segmented both have small paracotylar foramina (Supporting Information, File S1: +Fig. S2 +). + +The ribs are typical for alethinophidians, well curved, with a posterodorsal tubercle and a small foramen on the shaft just distal to the articular head. + + + \ No newline at end of file diff --git a/data/03/D0/58/03D0580CFFFDFF8CFF79FC0173CDE943.xml b/data/03/D0/58/03D0580CFFFDFF8CFF79FC0173CDE943.xml new file mode 100644 index 00000000000..cd6149afa31 --- /dev/null +++ b/data/03/D0/58/03D0580CFFFDFF8CFF79FC0173CDE943.xml @@ -0,0 +1,821 @@ + + + +A new fossil frog (Lissamphibia: Anura) from the Late Cretaceous of Brazil and the early evolution of neobatrachians + + + +Author + +Santos, Rodolfo Otávio +Instituto de Biociências, Universidade de São Paulo, São Paulo, Rua do Matão, Tv. 14, CEP 05508 - 090, São Paulo, SP, Brazil & Museu de Zoologia da Universidade de São Paulo, Avenida Nazaré 481, CEP 04263 - 000, São Paulo, SP, Brazil + + + +Author + +Carvalho, Alberto B. +Museu de Zoologia da Universidade de São Paulo, Avenida Nazaré 481, CEP 04263 - 000, São Paulo, SP, Brazil + + + +Author + +Zaher, Hussam +Museu de Zoologia da Universidade de São Paulo, Avenida Nazaré 481, CEP 04263 - 000, São Paulo, SP, Brazil + +text + + +Zoological Journal of the Linnean Society + + +2023 + +2023-11-15 + + +202 + + +1 + + +1 +29 + + + + +https://doi.org/10.1093/zoolinnean/zlad167 + +journal article +306819 +10.1093/zoolinnean/zlad167 +9cc8fa2d-6360-441d-9bb6-e186c471314c +0024-4082 +14542118 +1482C52-566F-4D11-8DFA-D6B20E2EB8A3 + + + + + + +Mariliabatrachus navai + +gen. et sp. nov. + + + + + + +urn:lsid:zoobank.org:act: +730D61E9-4502-4E7C-A07B-610BB48A637B + + + + + +Etymology: + +‘ +navai + +’, after the Brazilian palaeontologist William Roberto Nava, in recognition of his longstanding work discovering fossil vertebrates in the region of +Marília +municipality and vicinities. + + + + + + +Holotype +: + +MZSP-PV 25 +(a and b), housed at Museu de Zoologia da Universidade de São Paulo, represented by a nearly complete skeleton, comprising skull and postcranial remains, including vertebrae, scapular and pelvic girdles, and limb bones. + + + +Referred material: +MZSP-PV 26 (fragments of the skull, lower jaw, and postcranial remains, including pectoral girdle and the proximal portions of the forelimbs and anteromedial portions of vertebral column); MZSP-PV 27 (a fragmented braincase, including the otic capsules and parasphenoid); MZSP-PV 28 (an incomplete posterior portion of the skull, encompassing only the otic capsules); MZSP-PV 1326 (a partially preserved hindlimb, including tibiale, fibulare, tarsals, and the phalanges); MZSP-PV 1327 (partially preserved skull, including the left quadratojugal, and other indeterminate elements); MZSP-PV 1328 (elements of the pelvic girdle and distal part of the vertebral column); MZSP-PV 1329 (fragments of maxilla, scapula and sacral vertebra); MZSP-PV 1330 (an isolated fragment of a pterygoid); and MZSP-PV 1331 (isolated vertebrae). + + + + + +Diagnosis: +Mariliabatrachus navai + +gen. et sp. nov. +is a small anuran species that differs from pipoids by having well-developed mentomeckelian bones and a T-shaped parasphenoid, and shares with neobatrachians features like the presence of a neopalatine, eightprocoelouspresacralvertebrae, roundedsacraldiapophyses, and the lack of free ribs. The new species can be distinguished from other fossil and living neobatrachians by the presence of the following combination of characteristics (autapomorphies marked with an asterisk): azygous frontoparietal bearing ornamentations in form of shallow grooves and two laterally projected and rounded supraorbital flanges*; tectal roofing of +cavum cranii +composed by +taenia tecti transversalis +and +medialis +; cultriform process without ventral medial ridge and almost reaching the level of the +planum antorbitale +anteriorly; laterally expanded +crista parotica +; vomer present and bearing a straight tooth row; undivided sphenethmoid exposed between nasals and frontoparietal; +pars facialis +of the maxilla smooth, decreasing in height in the orbital region, and lacking anterodorsal and anteroventral processes; articulation between lower jaw and skull located anteriorly in relation to the occiput but laterally to the otic capsules; atlas with widely separated cotyles; omosternum present and mineralized; fusion between clavicula, scapula and coracoid; forked cleithrum; humerus with well-developed crista ventralis and ulnar epicondyle; intercalary elements absent; straight urostyle, ilium lacking supraacetabular fossa but presenting a low dorsal prominence and an elongated dorsal protuberance; acetabular fossa roughly cordiform, with a well-marked acetabular rim, except for its dorsalmost portion; ilial shaft bearing a moderately high dorsal crest; pubic region ossified; and femur nearly equal to tibiofibula in length and bearing a well-developed crest. + + + + +Type locality and horizon: +Outcrop located on the left margin of the Peixe river, known as Estrada Velha +, municipality of +Ocauçu +, +18 km +south-west of the municipality of Marília ( +49º56ʹ45ʹʹ W +, +22º20ʹ32ʹʹ S +). This deposit is traditionally assigned to the +Coniacian-Maastrichtian +( + +Castro +et al. +2018 + +) Adamantina Formation, part of the Bauru Basin. + + + + +Description + + +The hypodigm of + +Mariliabatrachus navai + +comprises 10 three-dimensionally preserved specimens ( +Fig. 2 +), encompassing almost all parts of the skeleton, except minor portions of the skull, forelimbs, and hindlimbs. The estimated snout–vent length of the +holotype +is +38.57 mm +. The presence of carpal elements, along with the ossification of the epiphyses on the long bones, and a well-ossified mentomeckelian bone, indicate that the +holotype +MZSP-PV 25 +( +Fig. 3 +) represents an adult individual. + + +Skull + + +Components of the skull are present in nearly all analysed specimens ( +holotype +and the +paratypes +MZSP-PV 26 +, +MZSP-PV 27 +, +MZSP-PV 28 +, +MZSP-PV 1327 +, +MZSP-PV 1329 +, and +MZSP-PV 1330 +), but are better preserved in the +holotype +. The skull of the +holotype +( +Fig. 4 +) is small and has approximately equal width and length. In dorsal view, it shows a roughly triangular outline (maximum width of +13.46 mm +, measured at the level of the lower jaw articulation; median length of +13.15 mm +, measured from the anterior margin of the premaxilla to the posterior margin of the occipital condyles). + + +Eoocranium +Both nasals are present in the +holotype +and arranged obliquely in relation to the sagittal plane of the skull. There is no medial contact between them, and they are positioned at the same level in relation to the anterior tip of the maxilla, but both conditions could not reflect the exact morphology of the fossil, as they could be also related to shifts that occurred during the fossilization process. Due to their damaged condition at the lateral margins, the exact morphology of this portion is uncertain. Their dorsal surface is smooth, without ornamentations. The nasals also bear pointed maxillary processes that project laterally, and the posterior margins of such processes contribute to the anterior margin of the orbits. The posterior portion of the nasal maxillary process abuts the maxilla, near the region in which the maxilla and the palatine articulate. There is no evidence of the contact among nasals with premaxilla, squamosal, or frontoparietal. + + + +Figure 2. +The hypodigm of + +Mariliabatrachus navai + +gen. et sp. nov +.. A, B, holotype MZSP-PV 25 (still articulated and almost complete skeleton). C, paratype MZSP-PV 26 (broken portions of skull, lower jaw fragments, and postcranial elements, including pectoral girdle, and parts of the forelimbs and vertebral column). D, paratype MZSP-PV 27 (fragment of a braincase). E, paratype MZSP-PV 28 (fragment of the posterior portion of the skull). F, paratype MZSP-PV 1326 (partially preserved hindlimb). G., paratype MZSP-PV 1327 (partially preserved skull). H, paratype MZSP-PV 1328 (elements of the pelvic girdle and distal column). I, paratype MZSP-PV 1329 (fragments of maxilla, scapula, and sacral vertebra). J, paratype MZSP-PV 1330 (isolated fragment of a pterygoid). K, paratype MZSP-PV 1331 (isolated vertebra). Scale bar = 1 cm. + + + + +Figure 3. +Details of MZSP-PV25, the holotype of + +Mariliabatrachus navai + +gen. et sp. nov +.. All the abbreviations are listed in the Appendix. Scale bar = 1 cm. + + + +The frontoparietals are fused into a single bone and lack a conspicuous suture line medially positioned. In dorsal view, the surface of this bone bears numerous irregular ornamentations, in the form of small and shallow grooves, more pronounced at its posteromedial portion. The frontoparietal covers anteriorly the posterior margins of the sphenethmoid. The fontanelle is absent, whereas short and posterolaterally directed supraorbital flanges are present on each lateral side of the frontoparietal. The +lamina perpendicularis +is present, being more pronounced at the posterior portion of the frontoparietal and narrowing anteriorly. As the posterolateral portions of the sphenethmoid were not preserved, it is not possible to determine if the +lamina perpendicularis +is fused to them or not. The posterior region of the frontoparietal is clearly fused to the otic capsules and the exoccipitals. The dorsal surface of the endocranium left an imprint on the ventral part of the frontoparietal (i.e. incrassatio frontoparietalis), comprising two faint but still noticeable fenestrae parietales (Supporting Information, +Fig. S5 +). + + + +Figure 4. +Skull of the holotype (MZSP-PV 25) of + +Mariliabatrachus navai + +gen. et sp. nov. +in dorsal (A), ventral (B), anterior (C), posterior (D), and right lateral (E), and left lateral (F) views. Scale bars = 2 mm. + + + +In the +holotype +, only the right premaxilla (Supporting Information, +Fig. S2A, B +) was preserved. This bone ( +2.11 mm +in length) is almost complete, lacking only the alary process and a portion of its lateral end. The +pars facialis +is smooth, without ornamentations. As only the base of the alary process was preserved, its size, orientation, and shape are uncertain. The +pars palatina +has a short and lingually projected palatine process, but due to its fragmentary condition, the configuration of its lateral end is uncertain. As the premaxilla is slightly shifted from its original position, the kind of contact with the maxilla is uncertain. The presence of teeth in the premaxilla could be inferred due to the presence of numerous pedicels, but the crowns were not preserved. + + +Only the left maxilla (total length of +8.04 mm +) was preserved in the +holotype +(Supporting Information, +Fig. S2G, H +). Its +pars facialis +has a wide orbital portion but narrows gradually posteriorly, and there are no ornamentations on its surface. In the anterior margin of the maxilla, the anterodorsal process is absent, whereas the anteroventral process, despite being present, is limited to a short projection. The kind of contact between the maxilla and quadratojugal (Supporting Information, +Fig. S3E, F +) is uncertain since the latter was preserved disarticulated from the rest of the skull. The +pars dentalis +bear pedicels along almost its entire length, being absent only at the last +3 mm +of its posterior portion. However, no complete teeth were preserved. Near the region of the maxillary preorbital process, the anterior branch of the pterygoid and the nasopalatine lateral tip are articulated, as well as the maxillary process of the nasals. + + +The triradiate squamosal is represented partially by the left element in the +holotype +and complete in +MZSP-PV 1327 +(Supporting Information, +Fig. S3C, D +), presenting a slender ‘T’-shape, without ornamentations on its dorsal surface. The squamosal is also slightly shifted from its original position but, based on its general shape, there is no contact between this bone and the azygous frontoparietal. The zygomatic ramus is short and narrow, and does not reach the maxilla. The otic ramus is short and narrow, slightly overlapping only the lateral portions of the +crista parotica +, whereas the ventral ramus is long, narrow, and straight. + + +In the +holotype +, both palatines [neopalatine of +Trueb (1993) +] were preserved (Supporting Information, +Fig. S2E, F +), being visible in both dorsal and ventral views. This bone is thin and positioned anterolaterally in relation to the sagittal line of the skull. Despite the lack of odontoids in the ventral surface of this bone, a short ventral ridge is present. The palatine is also partially covered by the sphenethmoid lateral process and by the maxillary process of the nasal. On its lateral margin, there is a contact with the +pars palatina +of the maxilla, near the preorbital process. + + +Vomers (Supporting Information, +Fig. S2C, D +) are present and bear teeth, which are positioned anteriorly in relation to the +planum antorbitale +. The tooth row is straight, but as both vomers are shifted from their original positions, it is difficult to determine if the vomerine tooth rows were arranged angled or perpendicular to the skull midline. An anterior process is present, but as both vomers and maxilla were shifted in the fossilization process, the contact between these elements is uncertain. The prechoanal process is elongated, whereas the postchoanal one, despite being broken, appears to be distinct but short. + + +In the +holotype +, the parasphenoid is partially preserved and exhibits a ‘T’-shape (i.e. the parasphenoid alae are perpendicular to the cultriform process), and it is not fused to other skull elements, as seen in the +paratype +MZSP-PV 1327 +(Supporting Information, +Fig. S1D, F, G +). The parasphenoid alae reach the otic capsules and their lateral margins are transverse and convergent medially. Poorly developed ventral crests are present on its ventral surface. The arrow-shaped and long cultriform process has parallel lateral margins and gradually narrows anteriorly. In its anteriormost portion, the cultriform process reaches the region of the +planum antorbitale +. There is no evidence of a longitudinal ridge on its ventral surface. + + +Both pterygoids were preserved in the +holotype +, but the right one is slightly shifted from its original position. The anterior ramus of the pterygoid (Supporting Information, +Fig. S3A, B +) is long, contacting medially the anteromedial portion of the +pars palatina +of the maxilla, but does not reach the +planum antorbitale. +There is no ventral flange on the anterior process, but the dorsal one is present, despite being weakly developed. The medial ramus is moderately developed and contacts the anterior face of the otic capsule. The posterior ramus is almost complete, lacking only its distal end. This process is short, not reaching the posterior limits of the skull. + + +Endocranium and stapes +The sphenethmoid is a single bone dorsally, with no evidence of ornamentations on its surface. In dorsal view, its posterior margins are partially covered by the frontoparietal. The anterior portion abuts the posterior margin of the nasals, but these margins do not overlap the sphenethmoid, which is exposed dorsally. Anteriorly, there is no bone extension of the sphenethmoid between the nasals. Laterally, the sphenethmoid bears an expansion that partially overlaps the palatine and forms part of the anterior margin of the orbits. The sphenethmoid is ossified only up to half of the +planum antorbitale +region, and the orbitonasal foramen is completely bound by bone (Supporting Information, +Fig. S1C +). In ventral view, this bone also forms a single element, bearing a smooth surface without conspicuous ornamentations. The lateral walls of the sphenethmoid, at the level of the optic foramen, are completely cartilaginous. The +septum nasi +is narrow and partially ossified, and the nasal capsules are medially close. + + +The prootics and the exoccipitals are fused into a single element, the otoccipital, which forms the otic capsule and the posteriorportionoftheskull.The +cristaparotica +isossifiedandhas a distal expansion. The deep scar located on the distal terminus of the +crista parotica +indicates the presence of a surface for articulation with the otic ramus of the squamosal. The exoccipitals are fused ventral and dorsally, surrounding completely the foramen magnum. The epiotic eminences are high, protrude posterolaterally, and are not covered by the frontoparietal. The occipital artery runs in an open groove located in the otoccipital. The occipital condyles are distinct and well separated from each other, with no confluence between them. + + +The stapes are well preserved in the +paratype +MZSP-PV 27 +and located beneath the otic capsules (Supporting Information, +Fig. S1E–G +). The preservation of a distinct stapes and the shape of the squamosal indicate the presence of a tympanic annulus. + + +Lower jaw + + +Elements of the lower jaw ( +Fig. 5 +) are better represented in the +holotype +( +11.38 mm +total length), but some bone fragments were also preserved in +MZSP-PV 26 +. The lower jaw of + +Mariliabatrachus navai + +is composed of well-ossified and edentate mentomeckelian bone, dentaries, and angulosplenials. In the +holotype +, only the mentomeckelians, right dentary, and both angulosplenials were preserved, whereas in +MZSP-PV 26 +only a fragmentary portion of the left angulosplenial is present. + + +The mentomeckelian bones are well ossified and fused to the dentaries ( +Fig. 5D, E +), despite the left one being broken and separated from the left dentary, which was not preserved. They are also slightly shifted from the region of the mandibular symphysis. There is no evidence of the presence of a lateral process on them. The right dentary of the +holotype +is slender, extending for approximately 66% of the total length of the lower jaw and forming most of the anterior portion of the lower jaw. Ornamentations are absent on its labial surface. There is no evidence of teeth or pedicels. The dentaryoverlapslaterallytheanteriorportionoftheangulosplenial, covering part of the Meckel’s canal. The angulosplenial is represented on both sides of the +holotype +, but the left element is better preserved. It is an elongated and arched bone, presenting a pronounced sulcus on its lateral face for the passage of the Meckel’s cartilage. The coronoid process is discrete and forms a relatively elongate, dorsomedially projected, and rounded lamina of bone in the posterior portion of the angulosplenial ( +Fig. 5C +). There is a lateral expansion of the angulosplenial at its posterior portion, whose dorsal face articulates with the jaw suspensorium. The articulation between the lower jaw and the skull is positioned anteriorly in relation to the occiput and laterally to the otic capsule. + + + +Figure 5. +Lower jaw of the holotype (MZSP-PV 25) of + +Mariliabatrachus navai + +gen. et sp. nov. +in dorsal (A), labial (B), and lingual (C) views. Details of the right dentary and mentomeckelian in labial (D) and lingual (E) views. All the abbreviations are listed in the Appendix. Scale bar = 1 mm. + + + +Postcranium + + +The postcranial bones of + +Mariliabatrachus navai + +are represented in the +holotype +and also in the +paratypes +( +MZSP-PV 26 +, +MZSP-PV 1326 +, +MZSP-PV 1328 +, +MZSP-PV 1329 +, and +MZSP-PV 1331 +), encompassing the vertebrae, pectoral and pelvic girdles, fore and hindlimbs. The only missing elements are the cartilaginous parts of the pectoral girdle and some phalanges. + + +Vertebral column +Despite its fragmentary condition, the vertebral column ( +Fig. 6A–C +) is complete (presacral vertebrae I to +VIII +, sacrum and urostyle) in the +holotype +, and represented by partially articulated presacral vertebrae ( +III +to +VII +) in the +paratype +MZSP-PV 26 +. In total, + +Mariliabatrachus navai + +presents eight presacral vertebrae, all of them procoelous, a condition observed mainly in the ventral view of the specimen +MZSP-PV 26 +. These vertebrae present neural arches not imbricated, with a short portion of the spinal canal exposed dorsally, and there is no evidence of the presence of a dorsal shield. + + +The first presacral vertebra (atlas) bears two well-defined and widely separated cotyles, which are positioned ventrolaterally in relation to the neural canal [type I of +Lynch (1971) +]. The atlas is articulated but not fused to the presacral +II +and does not exhibit a transverse process. The neural arch of the atlas has a low neural spine, whereas the postzygapophyses are limited to a short extension posteriorly directed. The presacral +II +is articulated to the atlas and exhibits elongated transverse processes projecting anterolaterally. The presacral +III +is articulated to the second presacral and also bears long transverse processes anterolaterally directed and distally expanded but lacking an uncinate process. The presacral +IV +is damaged in the +holotype +, and thus only disarticulated and indistinct portions of the neural arch and transverse processes were preserved. However, in +MZSP-PV 26 +the presacral +IV +is partially preserved, and the transverse processes are long and laterally directed. The presacral +V +is represented partially in the +holotype +and almost complete in +MZSP-PV 26 +, bearing transverse processes laterally directed. The presacral +VI +was preserved in the +holotype +and +MZSP-PV 26 +, despite being highly damaged in the +paratype +. This vertebra presents slender and long transverse processes laterally projected. The presacral +VII +, present in the +holotype +and +MZSP-PV 26 +, also bears transverse processes laterally projected, but they are considerably shorter than those of presacral +VI +. The following vertebra (presacral +VIII +) was preserved only in the +holotype +and exhibits short transverse processes (clearly shorter than the diapophyses of the sacral vertebra), which are projected slightly anterolaterally. The transverse processes of the vertebrae are unequal in length, according to the following sequence: +III +> +II +> +IV +≅ +V +≅ +VI +> +VIII +> +VII +. Neural spines are low in all presacral vertebrae. There is no evidence of the presence of free ribs associated with the transverse process of presacral vertebrae. + + +The sacral vertebra is almost entirely preserved in the +holotype +except for the distal ends of the diapophyses (total sacral length of ~ +6.59 mm +) ( +Fig. 6D–G +). However, in the specimen +MZSP-PV 1328 +(Supporting Information, +Fig. S4 +), the lateral margin of the sacral diapophysis is well preserved ( +1.16 mm +in length, +0.58 mm +in width). Two distinctive prezygapophyses are present and articulate the sacral vertebra with the postzygapophyses of the last presacral vertebra. This vertebra also exhibits two cylindrical but slightly flattened (ratio between height and length nearly equal to 0.5) sacral diapophyses posterolaterally directed, which are only slightly distally expanded. The articulation between this vertebra and the urostyle is bicondylar. The sacral diapophyses are oriented almost horizontally in relation to the horizontal plane in a posterior view. In +MZSP-PV 1328 +, an oval sesamoid is present on the distal margin of the sacral diapophysis. + + +The long urostyle of the +holotype +( +Fig. 6H–J +) is entirely preserved and is straight in lateral view, presenting a total length of +9.78 mm +. Next to the anterior margin, there is a foramen for the passage of the spinal nerve. The dorsal crest of the urostyle is present and well-developed, but it remains high only throughout its anterior half. Laterally positioned transverse processes are absent in the urostyle, whereas an anterodorsal process located on the anterior margin of the dorsal crest is present. + + +Pectoral girdle +The elements associated to the pectoral girdle of + +Mariliabatrachus navai + +( +Fig. 7 +) are present in the +holotype +and +MZSP-PV 25 +, but are better preserved in the +paratype +MZSP-PV 26 +. The pectoral girdle is arciferal and slightly shifted laterally in +MZSP-PV 26 +. The omosternum is present and mineralized (Supporting Information, +Fig. S6 +). The clavicle, coracoid, and scapula are fused. In +MZSP-PV 26 +, the clavicles are well preserved but are overlapping each other due to the shifting that occurred during the fossilization process. Both are strongly curved and their medial edges have no expansions, whereas their lateral margins are fused to the scapula and coracoid. Despite their shift, the clavicles are oriented anteromedially, and this condition is better observed in the right clavicle of +MZSP-PV 26 +. + + +The coracoids bear an expanded lateral margin fused anteriorly with the clavicle.These bones are arranged horizontally, presenting wide medial margins and an anterior edge slightly expanded anteriorly. The sternal ends are asymmetrical and only slightly expanded (less than half) in relation to the total length of the bone. The posterodorsal process is absent. Both scapulae (total length of +5.50 mm +) are present in MZSP-PV 26, but the right one is damaged and only its proximal margin is known, whereas the left one is more complete, showing a concave anterior margin, without processes or crests, and a roughly straight posterior margin. + + +The +pars acromialis +is well developed. The glenoid fossa (total length of +1.94 mm +) is completely ossified and closed by the fused clavicle lateral margins, coracoid, and proximal margins of the scapula. The scapula is relatively elongated, as the maximum width of the glenoid fossa is less than half of the total width of the scapular shaft. The medial cleft of the scapula is absent. The cleithra are only barely visible in the specimen +MZSP-PV 26 +, whereas in the +holotype +only a fragment assigned to the left cleithrum was preserved. This bone shows a straight lateral (scapular) margin and a bifurcated posterior margin, in which the anterior and posterior branches are separated by a distinct notch. +Forelimbs +The preserved components of the + +Mariliabatrachus navai + +are the humeri, radioulna, carpal elements, and some phalanges. Both humeri are preserved and complete in the +holotype +and +paratype +MZSP-PV 26 +. The humerus ( +Fig. 8A–D +) is elongated and presents a well-developed +crista ventralis +that reaches the median portion of the relatively narrow diaphysis. The +crista medialis +and the +crista lateralis +are absent. The humeral condyle is spherical, only slightly shifted laterally, well ossified, and large (its total diameter in relation to the total distal width at the epicondyle level is more than 0.58). The ulnar epicondyle is relatively well-developed, whereas the radial one is small. The cubital fossa is located immediately above the humeral condyle, being small and shallow. The radioulna ( +Fig. 8E, F +) is present in both limbs of the +holotype +. This bone bears a deep and concave olecranon fossa, and the olecranon process is well developed. Carpal bones (including pre- and postaxial elements) of both limbs were preserved in the +holotype +; however, due to shifts that occurred during the fossilization and their damaged condition, the following morphological assignments are tentative. Based on the positioning of the carpals 4 * 5 and the morphology of the metacarpals, we interpret that the left manus ( +Fig. 9 +) was completely flipped during fossilization. The carpal torsion is apparently present. Regarding the postaxial carpals, the ulnare and the distal carpal 3 are free (i.e. not fused), but the distal carpals 4 * 5 are fused into a single element, whereas the preaxial carpals (element Y and distal 2) are free. The morphology and the total number of the prepollical elements cannot be given, as only one element tentatively assigned to the prepollex was preserved. In the +holotype +, all the four metacarpals were preserved in the left manus but their distal margins are broken and the proximal ones are damaged, whereas in the right manus only two indeterminate metacarpals are still present, despite their highly damaged condition. As the left manus was flipped, the order of the metacarpals is reversed. The metacarpal +II +has a proximal margin ( +1.16 mm +of width) wider than the one of the metacarpal +III +( +0.88 mm +of width). The proximal margin of the metacarpal +III +exhibits only one articulation surface. In the right manus ( +Fig. 10 +), five elements were tentatively identified as the proximal and terminal phalanges. There is no evidence of the presence of intercalary elements between the only proximal and terminal phalanges still articulated. The terminal phalanges are conical and straight, presenting a rounded distal tip and a planar to convex proximal epiphysis. + + + +Figure 6. +Articulated vertebral column of the holotype (MZSP-PV 25) of + +Mariliabatrachus navai + +gen. et sp. nov. +in dorsal (A), ventral (B), and right lateral (C) views. Sacral vertebra in dorsal (D), ventral (E), anterior (F), and posterior (G) views. Urostyle in anterior (H), left lateral (I), and dorsal (J) views. All the abbreviations are listed in the Appendix. Scale bars = 1 mm. + + + + +Figure 7. +Pectoral girdle of the paratype (MZSP-PV 26) of + +Mariliabatrachus navai + +gen. et sp. nov. +in ventral (A) and dorsal (B) views. All the abbreviations are listed in the Appendix. Scale bar = 2 mm. + + + + +Figure 8. +Forelimb bones of the holotype (MZSP-PV 25) of + +Mariliabatrachus navai + +gen.et sp. nov +.. Left humerus in dorsal (A), medial + +(B), ventral (C), and lateral (D) views. Radioulna in ventral (E) and dorsal (F) views.All the abbreviations are listed in the Appendix.Scale bar = 1 mm. + + + +Figure 9. +Interpretative reconstitution of the left manus of the holotype (MZSP-PV 25) of + +Mariliabatrachus navai + +gen. et sp. nov. +in dorsal (A) and ventral (B) views. Left manus as preserved in the holotype in ventral (C) and dorsal (D) views. All the abbreviations are listed in the Appendix. Upper scale bar = 2 mm, lower scale bar = 1 mm. + + + + +Figure 10. +Right manus of the holotype (MZSP-PV 25) of + +Mariliabatrachus navai + +gen. et sp. nov. +in ventral (A), dorsal (B), and lateral (C) views. Details of the terminal and penultimate phalanges in lateral (D) and ventral (E) views. All the abbreviations are listed in the Appendix. Upper scale bar = 2 mm, lower scale bar = 1 mm. + + + + +Figure 11. +Pelvic girdle of the holotype (MZSP-PV 25) of + +Mariliabatrachus navai + +gen. et sp. nov. +in acetabular (A) and dorsal (B) views. All the abbreviations are listed in the Appendix. Scale bar = 2 mm. + + + +Pelvic girdle +Elements of the pelvic girdle ( +Fig. 11 +) are well preserved in the +holotype +and the +paratypes +MZSP-PV 1328 +and +MZSP-PV 1329 +. They are still articulated with the vertebral column and hindlimbs in the +holotype +, and there is only a faint suture line between the ilium, ischium, and pubis, and so they are fused. In acetabular view, the ilial shaft is moderately elongated (~3.5 times longer than the ilial body) and slightly arched dorsally. The ilial shaft bears a moderately high dorsal crest (~ +0.84 in +relation to the total high of the ilial shaft) extending almost throughout its entire length, except for the distal end. In medial view, there is a relatively deep and elongated sulcus between the dorsal portion of the shaft and the ventral portion of the dorsal crest. The cross-section of the ilial shaft is oval in anterior view, but the dorsal crest forms a thin projection dorsomedially oriented. In the ilial body, the dorsal prominence is present but low, being rounded and confluent with the dorsal crest of the ilial shaft, and also bears a laterally projected and elongated dorsal protuberance. The dorsal acetabular expansion is moderately developed ( +1.1 mm +in length), posterodorsally oriented, and positioned at the same height in relation to the dorsal prominence–protuberance complex. There is no supraacetabular fossa. The acetabular fossa is relatively shallow, wide ( +1.95 mm +length, +2.32 mm +width), and has a roughly cordiform outline, being located posteroventrally in relation to the dorsal prominence–protuberance complex. A well-pronounced acetabular rim surrounds almost entirely the margins of the fossa, except for its dorsalmost portion. The preacetabular zone is well-developed and leads to moderately wide ventral acetabular expansion ( +1.45 mm +in length) anteroventrally directed, slightly broader than the dorsal one. There is an angle of ~90º between the anterior margin of the ventral acetabular expansion and the ilial shaft. In ventral view, the surface of the ilial body is smooth, and there is no sign of a spiral groove. The ischium bears a well-developed posterodorsal region, which has a quadrangular outline, whereas its portion located posterior to the acetabular fossa is short and has a relatively straight margin. The portion of the pubis located below the acetabular fossa is short, except for the region next to the ventral acetabular expansion, which is elongated and forms a rounded projection oriented anteroventrally. + + +Hindlimbs +The elements that comprise the hindlimbs of + +Mariliabatrachus navai + +were preserved only in the +holotype +and the specimen +MZSP-PV 1326 +. Among the preserved bones in the +holotype +, there are the entirely preserved right femur ( +Fig. 12 +) and tibiofibular, and the partially preserved left femur and the right tibiale and fibulare. + + + +Figure 12. +Hindlimb bones of the holotype (MZSP-PV 25) of + +Mariliabatrachus navai + +gen. et sp. nov +.. Right femur in dorsal (A), lateral (B), and ventral (C) views. Right tibiofibula in lateral (D) view. All the abbreviations are listed in the Appendix. Scale bar = 1 mm. + + + + +Figure 13. +Tibiale, fibulare, and pes of the paratype (MZSP-PV 1326) of + +Mariliabatrachus navai + +gen. et sp. nov. +in ventral (A) and dorsal (B) views. All the abbreviations are listed in the Appendix. Scale bar = 1 mm. + + + +In MZSP-PV 1326, the left tibiale, fibulare, distal tarsals, metatarsals, prehallux, and some phalanges were preserved at least partially ( +Fig. 13 +). The remaining skeletal elements (i.e. the left tibiofibula and the right distalmost portions of the hindlimb) are absent. Both femora are completely ossified, exhibiting a roughly straight outline and a conspicuous femoral crest positioned along the proximal half of its length. The femur has a slightly short total length when compared with the tibiofibula ( +14.32 mm +and +15.59 mm +, respectively). The tibiofibula is nearly straight. Regarding the proximal tarsals ( +Fig. 13 +), the tibiale and fibulare (total length of +7.45 mm +), they are partially fused proximally (width at proximal epiphysis equal to +2.37 mm +) and distally (width at distal epiphysis equal to +2.48 mm +), and so there is a gap between their diaphyses. The distal tarsals 2 and 3 are fused, and another adjacent tarsal was tentatively interpreted as formed by the fusion of element Y and tarsal 1. Near to this element, there is a completely preserved element and a bone fragment tentatively identified as proximal and distal prehallical elements. Toes of the left pes were partially preserved in MZSP-PV 1326 ( +Fig. 13 +), including portions of metatarsals and proximal phalanges. Distal phalanges of the toes II, III, IV, and V are present but highly damaged. The only terminal phalanx present was tentatively associated with the toe II. This phalanx is roughly straight and its distal end is slightly knobbed. There is no evidence of the presence of intercalary elements. + + + + +Phylogenetic analyses + + +The morphology-based analysis under equal weights and unordered multistate characters resulted in 140 mostparsimonious trees (MPTs) of 1380 steps (CI = 0.172, RI = 0.554), the strict consensus of which is illustrated in +Figure 14 +. Bremer and jackknife support values for the nodes of the consensus are indicated at the base of each branch. Under these analytical conditions, the monophyly of Neobatrachia (i.e. a clade sister to + +Heleophryne + +) was not recovered. + +Mariliabatrachus navai + +was retrieved nested among a large clade of well-ossified hyloid neobatrachians (including both living and extinct species), although with weak support. In our analysis, the monophyly of this clade is supported by a single synapomorphy, Ch. 85: transverse processes/associated ribs of vertebra III lacking uncinate processes. A complete list with the common synapomorphies of all MPTs is available in the Supporting Information, +Fig. S13 +. + + +There are some differences between the topology obtained from this analysis and those generated under different analytical conditions (e.g. treating multistate characters as ordered or using implied weighting). Further details on these alternate analyses are provided in the Supporting Information, +Figs. S7–S +12 +. The strict consensus of the analysis using ordered multistate characters resulted in a large polytomy, with only a few clades emerging as monophyletic (e.g. a subgroup of ranoids and the species of + +Telmatobius + +). On the other hand, the other analyses (treating multistate characters as unordered but using implied weighting with different values of k) produced better-resolved topologies (e.g. Neobatrachia was recovered as monophyletic), although typically with low support. Additionally, neither Ranoidea nor Hyloidea have been retrieved as monophyletic. The phylogenetic positioning of + +Mariliabatrachus + +also varies depending on the analysis. In some cases, it was recovered as the sistertaxon of + +Uberabatrachus + +and + +Beelzebufo + +(k = 3), + +Eurycephalella + +(k = 7), or a group containing Late Cretaceous species and + +Calyptocephalella + +(k = 10; k = 20). + + + + \ No newline at end of file diff --git a/data/03/D0/58/03D0580CFFFDFF9DFEA4FDCF7057EEAE.xml b/data/03/D0/58/03D0580CFFFDFF9DFEA4FDCF7057EEAE.xml new file mode 100644 index 00000000000..c205e4690fb --- /dev/null +++ b/data/03/D0/58/03D0580CFFFDFF9DFEA4FDCF7057EEAE.xml @@ -0,0 +1,108 @@ + + + +A new fossil frog (Lissamphibia: Anura) from the Late Cretaceous of Brazil and the early evolution of neobatrachians + + + +Author + +Santos, Rodolfo Otávio +Instituto de Biociências, Universidade de São Paulo, São Paulo, Rua do Matão, Tv. 14, CEP 05508 - 090, São Paulo, SP, Brazil & Museu de Zoologia da Universidade de São Paulo, Avenida Nazaré 481, CEP 04263 - 000, São Paulo, SP, Brazil + + + +Author + +Carvalho, Alberto B. +Museu de Zoologia da Universidade de São Paulo, Avenida Nazaré 481, CEP 04263 - 000, São Paulo, SP, Brazil + + + +Author + +Zaher, Hussam +Museu de Zoologia da Universidade de São Paulo, Avenida Nazaré 481, CEP 04263 - 000, São Paulo, SP, Brazil + +text + + +Zoological Journal of the Linnean Society + + +2023 + +2023-11-15 + + +202 + + +1 + + +1 +29 + + + + +https://doi.org/10.1093/zoolinnean/zlad167 + +journal article +306819 +10.1093/zoolinnean/zlad167 +9cc8fa2d-6360-441d-9bb6-e186c471314c +0024-4082 +14542118 +1482C52-566F-4D11-8DFA-D6B20E2EB8A3 + + + + + + +Mariliabatrachus + +gen. nov. + + + + + + +urn:lsid:zoobank.org:act: +130B7020-BB71-4286-B26C-97354FA101F2 + + + + + +Etymology: +‘ + +Marilia + +’, after the municipality of +Marília +in the state of +São Paulo +, closest to the locality where the fossils were found, and ‘ +batrachus +’, derived from the Greek word +bátrakhos +, meaning frog. + + + + + +Type species: +Mariliabatrachus navai + +gen. et sp. nov +.. + + + + \ No newline at end of file