Caridina boehmei Klotz & Rintelen, 2013, n. sp.
Creators: name: Klotz, Werner
name: Rintelen, Kristina Von
Publisher: name: Zenodo
Year: 2013
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Malacostraca
Decapoda
Atyidae
Caridina
Caridina boehmei

description: Caridina boehmei n. sp.
(Figs. 1 A–Q, 2A–D, color plate 1A)
Material examined. Holotype male, cl 3.27 mm (MZB Cru 3179), Indonesia: Central Sulawesi, Tomori area, road Beteleme-Tomata, 02°5.972´S 121°9.672´E (Loc. 25-05, on mixed substrate), leg. T. von Rintelen, 12 Jan. 2005. Paratypes: 1 ovigerous female, cl 3.5 mm, 4 females, cl 3.0– 4.4 mm, 3 males, cl 2.7–3.2 mm (MZB Cru 3180), same data as holotype; 1 ovigerous female, cl 3.8 mm, 4 females cl 3.2–4.0 mm, 4 males cl 3.1–3.4 mm (ZMB 29308), same data as holotype. Others: 5 ovigerous females, cl 3.9–4.1 mm, 4 females, cl 3.0–4.0 mm, 3 males, cl 3.1–3.3 mm (MZB Cru 3181); 8 ovigerous females, cl 3.7–4.6 mm, 8 females, cl 3.3–4.4 mm, 7 males, cl 2.9–3.7 mm, 1 juvenile, cl 2.0 mm (ZMB 29449), Indonesia: Central Sulawesi, Tomori area, Puawu River, cascades at road Beteleme-Tomata, west of Wawopada, 02°1.314´S 121°12.042´E (Loc. 221-05, on roots), leg. T. von Rintelen, 21 Oct. 2005; several specimen from commercial aquarium import sold as “ Sulawesi Bee Shrimp” (OUMNH.ZC.2012-05-006, OUMNH.ZC.2012-05-007).
Comparative material examined. Caridina sulawesi Cai & Ng, 2009. Paratypes: 2 males, cl 3.30–3.75 mm, 2 females, cl 3.60–3.98 mm (ZRC), Sungei Beru, Kappang, Maros, Sulawesi Selatan, leg. P. Leclerc, 12 Aug.1990; 2 males, cl 2.95–3.18 mm, Indonesia: SW-Sulawesi, Bantaeng, near Loka village, 05°25.540'S 119°52.257'E, 2698 feet asl, leg. H. G. Evers, 21 Sept. 2010; 1 male, cl 4.08mm, 1 female, cl 5.12 mm, personal collection of the first author (59-10), Indonesia: South Sulawesi, Tilanga, 03°02.126'S 119°53.232'E, 2817 feet asl, leg. H. G. Evers, 24 Sept. 2010.
Caridina parvidentata J. Roux, 1904. Lectotype: ovigerous female, cl 4.0 mm (BNHM 8Ia), Malawa Quelle, Celebes (= Sulawesi), leg. G. D. Sarasin. Paralectotypes: 3 males, cl 2.60–2.93 mm, 2 females, cl 2.6–3.8 mm, 1 ovigerous female, cl 4.0 mm (BNHM 8Ia), Malawa Quelle, Celebes (5 Sulawesi), leg. G. D. Sarasin. Others: 4 males, cl 2.87–3.05 mm, 3 females, cl 2.91–4.03 mm (OUMNH.ZC.2012-05-008), Indonesia: SW-Sulawesi, Rantepao, tributary of Jalan Pongtiku, leg. R. Masche, 2005.
Description.
Cephalothorax and cephalic appendages. Carapace length 2.0– 4.6 mm (median 3.5 mm). Rostrum straight, short and pointed (Fig. 1 A), not reaching to end of basal segment of antennular peduncle mostly not reaching beyond length of eyes, 0.23–0.27 (median 0.25) times as long as carapace, unarmed or armed with 0–2 teeth on dorsal and ventral margin. Rostrum formula 0 + 0–2(0–1) / 0–2(0–1). Inferior orbital angle with a small antennal spine. Pterygostomian angle broadly rounded. Eyes well developed with globular cornea. Antennular peduncle 0.59–0.77 (median 0.64) times as long as carapace, first segment 1.97–2.2 (median 2.12) times as long as second segment, second segment 1.68–2.0 (median 1.95) times longer than third segment. Stylocerite reaching to 0.87– 0.96 (median 0.91) times of basal segment of antennular peduncle. Scaphocerite (Fig. 1 D) 2.88–3.40 (median 3.10) times as long as wide.
Abdominal somites, telson and uropods. Sixth abdominal somite 0.49–0.54 (median 0.53) times carapace length, 1.47–1.79 (median 1.63) times as long as fifth somite, 0.87–0.94 (median 0.90) times as long as telson. Telson length (Fig. 1 B, C) 2.90–3.02 (median 2.96) times as long as proximal wide, distal margin convex without median projection, with 3–4 pairs of dorsal spinules and one pair of dorsolateral spinules; distal end with 8–10 spines, sublateral pair distinctly shorter than lateral and inner spines. Preanal carina (Fig. 1 F) low, rounded, lacking a tooth or spine with few setae. Uropodal diaeresis (Fig. 1 E) with 15–16 movable spinules, outermost ones slightly longer than lateral angle.
Mouthparts and branchiae. Incisor process of mandible ending in irregular teeth, molar process truncated (Fig. 1 G). Lower lacinia of maxillula broadly rounded, upper lacinia elongate, with numerous distinct teeth on inner margin, palp slender with few simple setae at tip (Fig. 1 H). Upper endites of maxilla subdivided, palp slender, scaphognathite tapering posteriorly, fringed with long, curved setae at posterior margin (Fig. 1 I). Palp of first maxilliped ending in a fingerlike extension (Fig. 1 J). Podobranch on second maxilliped reduce to a lamina with few finger-like projections (Fig. 1 K). Third maxilliped with 2 arthrobranches, ultimate segment about as long as penultimate segment (Fig. 1 L). First pereiopod with an arthrobranch. Pleurobranchs present on all pereiopods. Well-developed (with hooks on distal end) epipods present on third maxilliped and first 2 pereiopods, lacking on pereiopods 3–5.
Pereiopods. Chela and carpus of first pereiopod stouter and broader than chela and carpus of second pereiopod (Fig. 1 M, N); chela of first pereiopod 2.10–2.32 (median 2.11) times as long as wide, 1.12–1.14 (median 1.13) times length of carpus; tips of fingers rounded, without hook; dactylus 1.20–1.38 (median 1.36) times as long as palm; carpus just slightly excavated distally 2.35–2.57 (median 2.40) times as long as wide, 1.27–1.34 (median 1.32) times length of merus. Merus 2.42–2.52 (median 2.48) times as long as wide, shorter than ischium. Chela of second pereiopod 2.78–3.06 (median 2.89) times as long as wide, 0.73–0.85 (median 0.76) times length of carpus; tips of fingers rounded, without hook, dactylus 1.15–1.32 (median 1.29) times as long as palm; carpus 4.81–5.91 (median 5.22) times as long as wide, 1.26–1.46 (median 1.39) times as long as merus; merus 4.22–4.65 (median 4.40) times as long as wide, shorter than ischium. Third pereiopod slender (Fig. 1 O–Q), not sexually dimorphic, dactylus 3.46–3.62 (median 3.50) times as long as wide (terminal claw and spines on flexor margin included), terminating in one large claw with five accessory spines on flexor margin; propodus 10.07–11.00 (median 10.67) times as long as wide, 3.40–3.60 (median 3.44) times as long as dactylus; carpus 5.00–5.47 (median 5.06) times as long as wide, 0.59–0.67 (median 0.60) times as long as propodus, 0.51–0.59 (median 0.55) times as long as merus; merus 6.73–7.71 (median 7.17) times as long as wide, 1.68–1.95 (median 1.81) times as long as carpus, bearing 3 small movable spinules on posterior margin of outer surface. Ischium with a small spinule. Fifth pereiopod slender (Fig. 2 A, B), dactylus 4.00–4.57 (median 4.12) times as long as wide (terminal claw and spines on flexor margin included), terminating in one large claw with 50–55 (28 in one male specimen) spinules on flexor margin, first spinule distinctly stronger than posterior ones; propodus 9.71–11.73 (median 10.42) times as long as wide, 2.47– 2.89 (median 2.61) times length of dactylus, carpus 4.29–5.11 (median 4.93) times as long as wide, 0.51–0.57 (median 0.54) times as long as propodus, 0.66–0.71 (median 0.68) times as long as merus; merus 6.48–7.00 (median 6.83) times as long as wide, 1.41–1.51 (median 1.48) times length of carpus, bearing one or two very small movable spinules on posterior margin of outer surface. Pleopods. Endopod of male first pleopod (Fig. 2 C) triangularly elongated, 2.28–2.44 times as long as proximal width (n=2), 0.38–0.39 times as long as exopod, without appendix interna. Appendix masculina on male second pleopod (Fig. 2 D) slender (6.91–8.11 times as long as distal wide), saccate, with some long spines on inner and numerous long spines distal margin, appendix interna reaching to about 0.8–0.9 of appendix masculina.
Reproductive biology. Ovigerous females with few eggs; size of eggs with developed 0.96–1.03 x 0.68–0.74 mm.
Colouration. Body semi-transparent. Three broad transverse blotches present on median part of the carapace, and on the third and the sixth abdominal segments. Remaining segments of abdomen and anterior part of the carapace with scattered white chromatophores.
Etymology. Caridina boehmei, new species, is named after Joachim Böhme for his kind support of our research on Sulawesi´s freshwater shrimp fauna.
Remarks. Caridina boehmei, n. sp., is similar to C. sulawesi Cai & Ng, 2009 showing a short and less strongly dentate rostrum and large-sized eggs. The new species can be distinguished from C. sulawesi by the shape and spinulation of the appendix masculina of the male second pleopod. This appendage appears saccate, i.e. 6.91–8.11 times as long as distal wide with numerous long spines on inner and distal margins, respectively in C. boehmei n. sp. vs. very slender (10.73–12.38 times as long as distal wide), stick-like and bearing few strong, short spinules arragned rougly in two rows along inner margin in C. sulawesi (Fig. 2 I). The endopod of male first pleopod is subtriangularly elongated, 2.28–2.44 times as long as wide and 0.38–0.39 times as long as endopod vs. sub-rectangular with distal margin angular, 2.87–3.27 times as long as wide and 0.47–0.65 times as long as exopod in C. sulawesi (Fig. 2 H). Further, the two species can be distinguished by the shape of the preanal carina (low vs. high in C. sulawesi), more slender carpus of first cheliped (2.35–2.57 times as long as wide vs. 1.51–2.06 times as long as wide in C. sulawesi) and by lacking sexual dimorphism of propodus of third pereiopod (with few spines arranged in 2 rows in males and females (Fig. 1 Q) vs. propodus of third pereiopods with numerous spines in males of C. sulawesi (Fig. 2 G)).
COLOUR PLATE 1. Life coloration of (A) Caridina boehmei n. sp. and (B) Caridina butonensis n. sp. (latter photograph courtesy of Hans Georg Evers).
The proposed new species can be differentiated from Caridina parvidentata J. Roux, 1904 by the more reduced epipodial apparatus (well developed epipods on first and second pereiopods vs. on first 3 pereiopods in C. parvidentata), the stouter scaphocerite (2.88–3.40 times as long as wide vs. 3.8 times in C. parvidentata), a more slender carpus on first cheliped (2.35–2.57 times as long as wide vs. 1.8 times as long as wide in C. parvidentata) and by the shape and spinulation of the appendix masculina (same differences as found in C. sulawesi).
The specimens collected from Puawu River show some differences compared to the type material: the rostrum is longer (reaching to the end of the first segment of the antennular peduncle, 0.29–0.40 times as long as carapace vs. not reaching to the end of the basal segment of the antennular peduncle, mostly not reaching beyond the eyes, 0.23–0.27 times as long as carapace), armed with more teeth (rostral formula 0+2–8(4–6)/0–5(1–3) vs. 0+0–2(0–1) /0–2(0–1)), the podobranch of the second maxilliped reduce to a simple lamina vs. reduce to a lamina with few finger-like projections, the movable spinules on the posterior margin of the merus of the third and the fourth pereiopods are well developed (vs. small) and the first cheliped with a somewhat shorter carpus (0.76–0.87 times as long as chela vs. 0.87–0.90 times as long in the type specimen). Although there are some morphological variations, however, the general morphological characters of these Puawu River specimens are agree well with those of the type specimens along with low genetic differences in a mDNA analysis (von Rintelen et al. 2010). Therefore, we tentatively assign these specimens to Caridina boehmei n. sp.
Distribution and habitat. Caridina boehmei n. sp. is only known from two small rivers in Central Sulawesi. At the type locality the shrimp was observed on rocks, which is a rather unusual substrate for riverine Caridina. The additional material was collected among roots near cascades of the Puawu River.
Caridina laroeha n. sp. (Figs. 3 A–L, 4A–H)
Material examined. Holotype male, cl 4.0 mm (MZB Cru 3770), Indonesia: Sulawesi, Laroeha, 02°28.226’S, 121°04.125’E, leg. F. Herder, 0 5 May 2004. Paratypes: 1 ovigerous female, cl 4.7 mm, 2 females, cl 4.0– 4.6 mm, 5 males cl 3.7–4.5 mm (MZB Cru 3771), same data as holotype; 1 ovigerous female, cl 4.2 mm, 1 females cl 4.6 mm, 6 males, cl 3.3–4.2 mm (ZMB 28093) same data as holotype. Others: numerous specimen (ZMB 28093) same data as holotype.
Comparative material examined. Caridina opaensis J. Roux, 1904. 2 females, cl 3.4 and 3.7 mm (ZMB 29338), Indonesia: Southeast Sulawesi, stream in Pruiala (Mulyocinta), N of Aopa area, 04°3.67’S 122°6.845’E, leg. M. Glaubrecht & K. & T. von Rintelen, 31 May 2005; 1 male, cl 3.1 mm, 3 females, cl 4.1–4.4 mm (ZMB 29008), Indonesia: Southeast Sulawesi, Benua River, northern arm, 04°13.388’S 122°6.397’E, leg. M. Glaubrecht & K. & T. von Rintelen, 31 May 2005; 3 males, cl 3.8–4.0 mm, 3 females, 4.1–4.5 mm (ZMB 29349), Indonesia: Southeast Sulawesi, Poniponiki, NW of Raterate, road Kendari-Kolaka, 04°2.115’S 121°52.816’E, leg. M. Glaubrecht & K. & T. von Rintelen, 30 May 2005. Caridina mahalona Cai, Wowor & Choy 2009. 3 males, cl 3.8– 4.0 mm (ZMB 29085), Indonesia: South Sulawesi, Larona catchment, Patingko River, road to Tabarano, off road Malili-Soroako N of Balambano, 02°38.222’S 121°14.56’E (Loc. 14-03), leg. K. & T. von Rintelen, 15 Sept. 2003; 1 male, cl 4.11 mm, 3 females, cl 4.0– 4.5 mm, 2 ovig. Females, cl 4.2–4.9 mm, Indonesia: South Sulawesi, at road Malili-Soroako, in Togo village, 02°36.53´S 121°15.56´E (loc. 34-03), leg. K. & T. von Rintelen, 15 Sept. 2003; 3 females, cl 4.2–5.3 mm (ZMB 29210), Indonesia: South Sulawesi, Towuti catchment, Bombongan, 02°52.38’S 121°22.54’E (Loc. 49-03), leg. K. & T. von Rintelen, 20 Sept. 2003.
Description. Cephalothorax and cephalic appendages. Carapace length 3.30–4.56 mm (median 3.82 mm). Rostrum (Fig. 3 A) long and slender, slightly sigmoid and upturned in distal part, reaching to or slightly overreaching distal end of scaphocerite, 0.8–1.2 (median 1.1) times as long as carapace, 0.4 distal dorsal margin unarmed, distal teeth of ventral margin more widely spaced. Rostrum formula 2–3 + 8–15 / 7–13. Antennal spine situated below inferior orbital angle. Pterygostomian angle rounded. Eyes well developed with globular cornea. Antennular peduncle 0.84–1.05 times as long as carapace in males, 0.70–0.83 times as long as carapace in females, first segment 1.45–1.68 (median 1.63) times as long as second segment, second segment 1.91–2.40 (median 2.18) times as long as third segment. Stylocerite reaching to, and 0.83–0.88 (median 0.88) times basal segment of antennular peduncle. Scaphocerite (Fig. 3 D) 3.42–4.08 (median 3.69) times as long as wide.
Abdominal somites, telson and uropods. Sixth abdominal somite 0.68–0.78 (median 0.75) times carapace length, 1.69–1.96 (median 1.74) times as long as fifth somite, 0.94–1.12 (median 0.98) times as long as telson. Telson length (Fig. 3 B, C) 3.07–3.25 (median 3.23) times as long as proximal wide, distal margin convex without median projection, with 4–6 pairs of dorsal spinules and one pair of dorsolateral spinules; distal end with 7–9 rather short spines, lateral pair longer than inner spines. Preanal carina (Fig. 3 F) with a distinct hook. Uropodal diaeresis (Fig. 3 E) with 9–12 movable spinules, outermost one shorter than lateral angle.
Mouthparts and branchiae. Incisor process of mandible ending in irregular teeth, molar process truncated (Fig. 3 G). Lower lacinia of maxillula broadly rounded, upper lacinia elongate, with numerous distinct teeth on inner margin, palp slender with few simple setae (Fig. 3 H). Upper endites of maxilla subdivided palp slender, scaphognathite tapering posteriorly, fringed with long, curved setae at posterior margin (Fig. 3 I). Palp of first maxilliped ending in a broad triangular extension (Fig. 3 J). Podobranch on second maxilliped reduce to a lamina (Fig. 3 K). Third maxilliped with 2 arthrobranches, ultimate segment about as long as penultimate segment (Fig.
3L). First pereiopod with an arthrobranch, pleurobranchs present on all pereiopods. Well developed (with hooks on distal end) epipods present on third maxilliped and first 2 pereiopods, reduced epipod (short, without distal hook) present on third pereiopod and epipod absent on pereiopods 4 and 5.
Pereiopods. Chela and carpus of first pereiopod stouter and broader than chela and carpus of second pereiopod (Fig. 4 A, B); chela of first pereiopod 2.19–2.32 (median 2.23) times as long as wide, 1.20–1.34 (median 1.30) times length of carpus; tips of fingers rounded, without hook; dactylus 0.97–1.24 (median1.20) times as long as palm; carpus scarcely excavated distally, 2.00–2.66 (median 2.26) times as long as wide, 1.09–1.20 (median 1.14) times length of merus. Merus 2.42–2.84 (median 2.68) times as long as wide, as long as ischium. Chela of second pereiopod 2.83–3.09 (median 3.04) times as long as wide, 0.78–0.85 (median 0.79) times length of carpus; tips of fingers rounded, without hook, dactylus 1.18–1.57 (median 1.39) times as long as palm; carpus 4.83–5.73 (median 5.24) times as long as wide, 1.30–1.45 (median 1.40) times as long as merus; merus 4.42–4.76 (median 4.48) times as long as wide, as long as ischium. Third pereiopod slender (Fig. 4 C, D), not sexually dimorphic, dactylus 3.94– 4.29 (median 4.16) times as long as wide (terminal claw and spines on flexor margin included), terminating in one large claw with 6–9 accessory spines on flexor margin; propodus 11.61–14.00 (median 12.41) times as long as wide, 3.41–3.92 (median 3.57) times as long as dactylus; carpus 5.44–5.62 (median 5.53) times as long as wide, 0.56–0.63 (median 0.60) times as long as propodus, 0.46–0.55 (median 0.51) times as long as merus; merus 8.77– 9.17 (median 8.88) times as long as wide, 1.82–2.18 (median 1.99) times as long as carpus, bearing 2 or 3 movable spinules on posterior margin of outer surface. Ischium with a small spinule. Fifth pereiopod slender (Fig. 4 E, F), dactylus 3.75–5.20 (median 4.84) times as long as wide (terminal claw and spines on flexor margin included), terminating in one large claw with 33–60 spinules on flexor margin, propodus 11.45–13.00 (median 12.52) times as long as wide, 2.80–3.65 (median 2.97) times length of dactylus, carpus 4.60–6.28 (median 4.88) times as long as wide, 0.48–0.54 (median 0.50) times as long as propodus, 0.57–0.61 (median 0.61) times as long as merus; merus 7.04–9.35 (median 7.76) times as long as wide, 1.63–1.77 (median 1.65) times length of carpus, bearing 3 or 4 movable spinules on posterior margin of outer surface. Pleopods. Endopod of male first pleopod (Fig. 4 G) subtriangular, long setae fringe along outer and distal margins, few medium-length setae fringe along inner margin, 2.07–2.18 times as long as proximal width (n=2), 0.26–0.27 times as long as exopod, without appendix interna. Appendix masculina on male second pleopod (Fig. 4 H) slender, 7.8 times as long as distal width, rod-shaped, with some long spines on inner and distal margins, appendix interna 0.85 as long as appendix masculina.
Reproductive biology. Ovigerous females with few eggs; size of eggs (from one female with undeveloped eggs) 0.94–0.99 x 0.57–0.60 mm.
Colouration. Unknown.
Etymology. The new species is named after its type locality – the Laroeha River, Sulawesi, Indonesia.
Remarks. With its slender rostrum and unarmed distorostral dorsal margin, Caridina laroeha n. sp. resembles C. opaensis and C. mahalona. Caridina laroeha n. sp. differs also from C. opaensis by having a longer rostrum (reaching to or slightly overreaching distal end of scaphocerite vs. mostly reaching to or slightly overreaching third segment of antennular peduncle in C. opaensis) with less postorbital teeth (2–3 vs. 3–7 in C. opaensis) but more ventral teeth (7–13 vs. 3–6 in C. opaensis) and 0.25 distal dorsal margin unarmed (vs. less than 0.25 in C. opaensis). Penultimate segment of third maxilliped as long as or slightly longer than ultimate vs. much longer in C. opaensis). The sixth abdominal segment is also longer (0.68–0.78 times carapace length vs. 0.53–0.63 times carapace length in C. opaensis), the number of the spinule on uropodal diaeresis is lower (9–11 vs. 12–19 in C. opaensis), the number of the spinule on the dorsal margin of the telson is higher (4–6 pairs vs. 3–4 pairs in C. opaensis), and the lateral spines on the distal end of the telson are longer than the inner spines (vs. as long as the inner spines with the second pair shorter in C. opaensis). However, 2 out of 6 of the comparative specimens of C. opaensis from Poniponiki (ZMB 29349) have relatively longer rostrum which are reaching to the end of the scaphocerite. Thus, these specimens resemble C. laroeha n. sp. in shape and length of the rostrum. Other characters of these 2 specimens of C. opaensis from Poniponiki (e.g. number of postorbital teeth and spinules on uropodal diaeresis, relative length of segments of third maxilliped 6. abdominal segement and number and length of spines on telson) resembles typical specimens of C. sulawesi.
Caridina laroeha n. sp. differs also from C. mahalona by having a longer rostrum (0.82–1.20 times carapace length vs. 0.52–0.75 carapace length in C. mahalona), less number of ventral rostral teeth (8–13 vs. 0–7 in C. mahalona), shorter stylocerite (0.83–0.88 as long as the first segment of antennular peduncle vs. 0.9–1.0 as long as the first segment of antennular peduncle in C. mahalona), relatively shorter penultimate segment of third maxilliped (as long as or slightly longer than ultimate vs. much longer penultimate in C. mahalona), epipod well developed on the second pereiopod but reduced on the third pereiopod (vs. absent from the third to the fifth pereiopods in C. mahalona), the carpus of the first and the second chelipeds are stouter (2.00–2.66 and 4.83–5.73 times as long as wide vs. 2.75–3.44 and 5.39–7.07 times as long as wide, respectivelyin C. mahalona), the carpus of the first pereiopod is relatively short (1.09–1.20 as long as merus vs. 1.33–1.44 as long as merus in C. mahalona), more spines on the dactylus of the third pereiopod (6–9 vs. 5–6 in C. mahalona), more slender sixth abdominal segment (0.68–0.78 times as long as carapace vs. 0.54–0.61 times as long as carapace in C. mahalona), less number of spinule on the uropodal diaeresis (9–12 vs. 12–15 in C. mahalona) and longer disto-lateral spines of telson compare to the inner spines (vs. as long as the inner spines with second pair shorter in C. mahalona).
Distribution and habitat. Caridina laroeha n. sp. is only known from the type locality—the Laroeha River in the western part of Central Sulawesi.
description: Published as part of Klotz, Werner & Rintelen, Kristina Von, 2013, Three new species of Caridina (Decapoda: Atyidae) from Central Sulawesi and Buton Island, Indonesia, and a checklist of the islands' endemic species, pp. 554-570 in Zootaxa 3664 (4) on pages 555-564, DOI: 10.11646/zootaxa.3664.4.8, http://zenodo.org/record/215621