Gastrotheca lojana Parker 1932
Creators: name: Carvajal-Endara, Sofía
name: Coloma, Luis A.
name: Morales-Mite, Manuel A.
name: Guayasamin, Juan M.
name: Székely, Paul
name: Duellman, William E.
Publisher: name: Zenodo
Year: 2019
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Anura
Hemiphractidae
Gastrotheca
Gastrotheca lojana

description: Gastrotheca lojana Parker 1932
Gastrotheca marsupiata lojana Parker, 1932:25.— Holotype: BM 1947.2.31.13 (Fig. 25B) from Loja, Loja Province, Ecuador. Gastrotheca monticola (part)— Duellman & Hillis 1987:158.
Gastrotheca (Duellmania) lojana — Dubois, 1987:33.
Referred specimens. Ecuador: Azuay: CJ 390, 401–2, from Oña, 2272 m (03° 27' 41.04" S, 79° 09' 46.47" W), on 15 June 2011 by Elicio E. Tapia, Sofía Carvajal-Endara and Henry Grefa; KU 138401–3, from Girón, 2310 m (03° 10' 00.12" S, 79° 07' 59.88" W), on 0 6 June 1970 by Thomas A. Fritts; QCAZ 2692, from Oña, 2272 m (03° 27' 41.04" S, 79° 09' 46.47" W), on 1 December 1999 by Luis A. Coloma and Luis E. López; QCAZ 26314–5, 26318, 26322–3, 26327–8, 26334–5, 26337–8, from Oña, 2272 m (03° 27' 40.43" S, 79° 09' 45.11" W), on 8 August 2003 by Luis A. Coloma and Ítalo G. Tapia, QCAZ 31521–3, from Oña, 2272 m (03° 27' 41.04" S, 79° 09' 46.47" W), on 27 December 2005 by Ítalo G. Tapia and Giovanni Onore; QCAZ 32212, from Oña, 2272 m (03° 27' 41.04" S, 79° 09' 46.47" W), on 2006 by Luis A. Coloma; QCAZ 32571, from Oña, 2272 m (03° 27' 40.43" S, 79° 09' 45.11" W), on 8 August 2003 by Luis A. Coloma, Erik R. Wild, Andrés Merino-Viteri, Ítalo G. Tapia and Edwin Patricio Vargas; QCAZ 42725, from San Fernando, Laguna de Busa, 2834 m (03° 09' 15.80" S, 79° 15' 49.03" W), on 22 October 2007 by Sofía Carvajal-Endara, Amaranta Carvajal-Campos, Andrea Carvajal Endara. Loja: BM 1947.2.31.6–10, BM 1947.2.31.13–18, from Loja, 2150 m (04° 00' 00" S, 79° 13' 00.12" W), on 1930–1 by Clodoveo Carrión Mora. KU 120673 –4, from Loja, 2150 m (04° 00' 00" S, 79° 13' 00.12" W), on 9 June 1968 by John D, Lynch; KU 120675, from 2 km E Loja, 2100 m (03° 59' 56.4" S, 79° 13' 00.12" W), on 9 June 1968 by John D, Lynch, KU 138233, from 3 km W Loja, 2150 m (03° 58' 47.99" S, 79° 12' 54" W), on 21 June 1970 by Thomas H. Fritts; KU 138234, from 10 km W Loja, 2500 m (03° 59' 56.4" S, 70° 14' 43.08" W), on 27 June 1970 by Thomas H. Fritts; KU 138235–6, from 5 km N Loja, 2150 m (03° 52' 00.12" S, 79° 13' 00.12" W), on 28 June 1970 by Thomas H. Fritts; KU 138404–9, from Saraguro, 2412 m (03° 36' 00" S, 79° 13' 00.12" W), on 19–20 June 1970 by Thomas H. Fritts; KU 148568, from Saraguro, 2412 m (03° 36' 00" S, 79° 13' 00.12" W), on 22 May 1971 by Richard M. Montanucci; KU 178482–95, from 2 km SSW Saraguro, 2569 m (03° 38' 22.92" S, 79° 13' 59.87" W), on 5 January 1978 by John D. Lynch; KU 178496–7, from 2.1 km N Saraguro, 2575 m (03° 36' 41.76" S, 79° 13' 00.12" W), on 7 January 1978 by John D. Lynch; QCAZ 22371, from Loja Zamora Huayco, 3018 m (04° 05' 49.9806" S, 79° 10' 02.499" W), on 12 December 2002 by Diego Almeida-Reinoso; QCAZ 30788, from Saraguro, 2412 m (03° 36' 00" S, 79° 13' 00.01" W), on 20 September 1978 by MC; QCAZ 34505, from 2 km SW Saraguro, 2569 m (03° 38' 22.99" S, 79° 14' 24" W), on 5 January 1978 by John D. Lynch. El Oro: QCAZ 32724, from Guanazán, 2984 m (03° 27' 18.43" S, 79° 29' 23.64" W), on 3 December 2006 by Silvia Aldás Alarcón.
Diagnosis. Included in the genus Gastrotheca by having a closed brood pouch on dorsum of female. A moderately large species (54.1–76.1 mm SVL in females, n = 12; 40.2–61.0 mm SVL in males; n = 24) with tibia length 41¯56% SVL, longer than foot; (2) interorbital distance about 1.6 times width of upper eyelid; (3) skin on dorsum finely granular, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus distinct, smooth; (7) Fingers I and II about equal in length, discs on fingers about twice width of digits proximal to discs; (8) fingers unwebbed; (9) webbing between external toes extending to the antepenultimate subarticular tubercle on Toe IV, to point midway between penultimate and distal subarticular tubercles on Toe V; (10) in life, dorsum green, brown, or tan with dark paravertebral marks; (11) markings on the head consisting of pale labial stripe broader at the posterior half of lip and dark interorbital bar or two blotches connected or not to paravertebral marks; (12) pale cream or white dorsolateral stripe fragmented; (13) flanks dark brown, green or tan with pale cream spots ventrally, anterior and posterior surfaces of thighs, groin, and dorsal surface of foot with contrasting pattern of heavily white mottling on a dark background, with bluish or greenish tinges; dorsal surfaces of limbs with dark brown or green bars; (14) venter creamy white with dark brown dots or marks; (15) brood pouch single, dorsal.
Gastrotheca lojana most closely resembles three other species in southern Ecuador (G. elicioi, G. cuencana, and G. litonedis) and one species in northern Peru (G. monticola). Gastrotheca lojana differs from G. elicioi in some coloration patterns (described above; also, compare in Figs. 10L, 29, 30 vs 12–14). A black canthal stripe is absent in G. lojana, but present in G. elicioi. The anterior and posterior surfaces of thighs and groin are heavily mottled in G. lojana, but slightly mottled in G. elicioi. The interorbital mark in G. lojana is a transverse bar that sometimes is divided into two blotches and that is usually connected with two broad paravertebral marks. In G. elicioi dorsal marks, when present, consist of a triangular interorbital mark, which may or may not be connected to
two narrow and curved paravertebral marks. Gastrotheca lojana has a conspicuous, elevated row of dorsolateral warts, whereas it is barely raised in G. elicioi. In addition, in G. elicioi the dark bars on limbs, when present, are shorter and less defined than in G. lojana. Gastrotheca lojana differs from its most related species, G. cuencana and G. litonedis (compare Figs. 10L, 29, 30 vs 2–4, 26–27) by having: a cream venter with dark flecks or spots (uniform creamy white in G. cuencana and pale brownish gray in G. litonedis); an interorbital bar or blotches usually connected to paravertebral marks (absent in G. cuencana and G. litonedis); dorsal surfaces of limbs with dark transverse bars (irregular dark blotches in G. cuencana and usually unmarked in G. litonedis); anterior and posterior surfaces of thighs and groin heavily mottled (translucent cream without marks in G. cuencana and usually brown and slightly molted in some G. litonedis); dorsal surfaces of fingers cream or with dark brown spots (cream in G. cuencana and of same color of the rest of the body in G. litonedis); a fragmented dorsolateral stripe (continuous in G. cuencana and G. litonedis); and complex call (simple call in G. cuencana and G. litonedis). The genetic divergence between G. lojana and G. litonedis is 2.5% and between G. lojana and G. cuencana is 2.8% (in a DNA dataset of 438 bp, 16S gene). Gastrotheca lojana differs from G. monticola by having the concealed surfaces of thighs, groin and dorsal surfaces of the feet with white mottling on a dark background, whereas in G. monticola the axilla and groin, concealed surfaces of thighs, and dorsal surfaces of the feet are green with black spots on a green or tan background.
A distinctly different species, G. pseustes, occurs sympatrically with G. lojana in part of its range. Gastrotheca lojana differs from G. pseustes by having finely granular skin on the dorsum (smooth to coarsely granular in G. pseustes), larger digital discs, and pale dorsolateral and supracloacal stripes. Also the call of G. lojana can be easily distinguished form the call of G. pseustes by the amplitude modulation of the longer note, shorter call duration, higher short note rate, shorter interval between notes, fewer pulses, lower pulse rate, a much lower dominant frequency, and a lower 90% bandwidth frequency.
Variation. Morphometric variation of 12 females and 24 males is summarized in Table 3. Females are larger than males (61.6± 5.7 mm; 49.4± 4.5 mm). All adults have a moderate supratympanic fold, which usually extends from the superior part of tympanum to point just above the insertion of forelimb. The dorsolateral row of warts can be less conspicuous in some specimens. Each vomer has four to ten teeth (5.7±1.4, n = 44).
Color variation in preservative. In most preserved specimens the dorsum is bluish gray or brown with darker interorbital, paravertebral marks and bars on the limbs. Some specimens also have dark flecks densely distributed on dorsum. All specimens lack a dark canthal stripe and have a cream white supralabial stripe, which is broader at the posterior margin of the lip and that is extended from the posterior margin of the lip to the insertion of the forelimb. Most of the specimens have a dark brown stippling on the supralabial stripe. A fragmented supracloacal and heel stripes are present; also a white mark extends laterally from the vent. The flanks are dark gray with black and white dots; the posterior part of the flanks, groin, concealed surfaces of thighs and dorsal surface of foot are heavily mottled. The venter is white and varies from having a few dark flecks to having a heavily mottled pattern.
Color variation in life. (Figs. 10L, 29, 30). In living individuals the color of the dorsum is green (CJ 408), brown (CJ 405) or tan (CJ 407) with darker interorbital and paravertebral marks. The interorbital mark is usually a bar that connects the two eyelids; however, in some individuals it is divided into two blotches (CJ 411). Paravertebral marks are broad and are usually connected with the interorbital mark (CJ 407) or, in some individuals (CJ 405), they are fused at the level of the scapular region. The venter is white in most specimens, some have dark flecks and spots evenly distributed, and others have a heavily mottled pattern. On the head there is a cream supralabial stripe, which is broader at the posterior margin of the lip and extends from the posterior margin of the lip to the insertion of the forelimb. The tympanum is brown, tan or olive green. The iris is reddish bronze with a few black reticulations. The flanks are brown, tan, or green with black and white spots. The posterior part of the flanks, groin, anterior and posterior surfaces of thighs have a heavily mottled pattern, some individuals have green or purple tinge on the groin and posterior surface of the thighs (Fig. 30). Supracloacal and heel stripes are cream or tan. The ventral surfaces of the shanks have a faint tinge of pale blue.
Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 36, from a series (CJ 1948) obtained from a pond at Oña, 2272 m, Azuay Province, Ecuador, by Elicio E. Tapia, Sofía Carvajal-Endara, and Henry Grefa on 15 June 2011 (Fig. 5C). Total length 55.7; body length 19.8 (36% of total length). Body ovoid in dorsal and lateral views, slightly depressed; throat shape concave in lateral profile; body width at the level of spiracle 12.3, and height at same position10.2; head width at level of eyes 10.5. Lateralline system barely visible, supraorbital and infraorbital lines both originating at tip of snout, running in parallel to the eye and making contact immediately behind the eye; angular line descending vertically from just posterior of eye to throat; anterior oral line descending vertically from level of oral disc and just anterior to level of nares to throat there it curve to parallel infraorbital line, thereby forming a continuous circuit with angular and loreal lines; dorsal body and middle body lines present.
Nostril medium sized (in proportion to body length), ovoid, protruding, with a fleshy annulus, its opening directed anterolaterally. Snout–nostril distance 2.8; internarial distance 3.2. Eye positioned and directed dorsally, not visible from below; eye length 2.1, eye width 1.9; interorbital distance 5.7. Spiracle sinistral, located ventrolaterally; spiracular opening directed posteriorly; distance from tip of snout to spiracular opening 13.7; spiracle end rounded, attached to body wall, inner wall of spiracle not evident; tube length 4.0, tube transverse width 2.9. Vent tube dextral, its opening directed posteriorly, tube length 3.7, tube transverse width 2.1. Tail length 37.0, caudal musculature slender, narrowing gradually until tail terminus; caudal muscle height 5.2, caudal muscle width 4.3; caudal fins well developed and about the same size, originating near tail-body junction; maximum height of tail 13.6; tail terminus rounded, caudal musculature not reaching fin terminus.
Oral disc small, ventral, located near tip of snout, not protruding laterally beyond body, not visible dorsally; transverse width 4.1. It is surrounded by a uniserial row of short, marginal papillae, interrupted medially on upper lip; lower lip papillae alternating in and out, giving the appearance of two series; upper lip with 21 papillae on right side and 19 papillae on left side; lower lip with 63 marginal papillae; upper jaw sheath medium-sized, forming a finely serrated, smooth arch, height 0.4, transverse width 3.0 (48% of oral disc width); lower jaw sheath V- shaped, open and finely serrated, width 2.1, height 0.7. Labial tooth row formula 2/3(1), tooth rows lengths: A1: 4.2, A2: 4.1, P1 right row 1.9, P1 left row 1.6, P1 gap 0.3, P2: 3.2, P3: 3.5. (Fig. 6C).
Color in preservative. Dorsum dull gray, with paler (translucent) areas on the flanks and snout. Caudal muscles pale gray with small cream flecks; dorsal and ventral fins translucent with evenly distributed cream flecks, except at tip where flecks are absent. Venter translucent on throat and belly regions, guts not exposed, white pigment partially covers the belly, producing a black and white mottling; eyes lavender gray with white flecks; oral apparatus translucent.
Color in life. In dorsal and lateral views, body olive-cream; loreal and snout areas palest. Ventrally, guts not visible, belly white with gray marks; gills visible through the throat, with a reddish hue. Caudal musculature cream-gray; proximal third reddish with myomeres and medial line well defined; distal two thirds with small cream flecks; dorsal and ventral fins translucent, suffused with minute cream flecks, clustering in rounded patches near borders of fins; most dense near tail-body junction. Spiracle, oral apparatus, and legs olive-cream. Iris gold.
Variation. Variation of 28 meristic characters of tadpoles in Stages 34–41 (CJ 1948) are shown in Table 9. Total length varies between 40.9 (Stage 34) and 61.2 (Stage 41) and tail length proportion varies from 59.1 to 63.7 until stage 41; labial tooth row formula 2/3(1). Number of marginal papillae varies among specimens and Gosner stages, variation in number of ventral papillae at lower lip is moderate (60 and 69). The color of venter varies from nearly plain gray (CJ 4303) (Fig. 31) to gray flecks in a white background (CJ 1948) (Fig. 5C). Duellman (2015) described a tadpole (KU 203548) from 7.9 Km W Loja. Nonetheless, it may also belong to either G. pseustes, G. psychrophila, or G. turnerorum.
We documented changes during ontogenetic development of CJ 4303, 4304 (Figs. 31 A–C, 32). At Stage 46, the dorsum is nearly uniform clear brown; the paraverterbal marks and interorbital marks are green and vary from nearly absent to well defined.
Comparisons. Tadpoles of Gastrotheca lojana may occur in sympatry with those of G. elicioi, G. psychrophila (tadpole unknown, see Remarks under G. elicioi tadpole account), G. pseustes 1, and G. turnerorum in the Loja- Abra de Zamora region, with G. litonedis and G. pseustes 2 (tadpole not described), in the Laguna de Busa area and with at least G. pseustes in other localities (compare in Fig. 5). Gastrotheca lojana differs from G. elicioi by lacking a dorsal gray-pigmented fin that abruptly arises from the body, from G. pseustes 1 by lacking a reticulated pattern on flanks, from G. turnerorum by having a more pointed tail terminus and lacking bold cream marks in a dorsal line of caudal musculature, and from G. litonedis by having a highest tail dorsal fin.
Vocalization. Five individuals of Gastrotheca lojana were recorded from one location in Azuay Province (one individual from San Fernando, Laguna de Busa) and from one location in Loja Province (four individuals from Oña) (Appendix III). The advertisement call of G. lojana is a complex call, composed of one long pulsed note and followed (or not) by one or two short, single-pulsed notes (Fig. 17 H–N). The long note had a mean duration of 0.562 s (SD = 0.125) and consisted on average of 17.59 (SD = 2.418) distinct pulses (pulse series), separated by silent intervals (amplitude modulation of 100%). The amplitude of the long note increases gradually towards the end, without falling. The short notes have a mean duration of 0.064 s (SD = 0.027) and the inter-note interval is on average of 0.356 s (SD = 0.107). The mean dominant frequency is 1118.1 Hz (SD = 126.872), with a mean 90% bandwidth of 986.1–1348.4 Hz. The fundamental frequency is not clearly recognizable; when visible, 5 to 7 harmonics are distinguishable in the short notes.
Comparisons. The advertisement call of Gastrotheca lojana is most similar to that of G. testudinea, but G. lojana has a shorter call duration, higher short note rate, shorter long notes, short notes duration, shorter inter-note intervals, higher pulse rate, a higher dominant frequency and a higher 90% bandwidth frequency compared with the call of G. testudinea (Table 5). The call of G. lojana can be easily distinguished form the calls of G. yacuri and G. pseustes by the amplitude modulation of the longer note, shorter call duration, higher short note rate, shorter inter-note interval, smaller number of pulses, lower pulse rate, a much lower dominant frequency, and a lower 90% bandwidth frequency. Also, G. lojana emits only one long note, whereas G. yacuri emits up to three (usually 2) long notes per call and G. pseustes emits a larger number of long and short notes (Table 5).
Distribution and ecology. Gastrotheca lojana is known in basins within Azuay, Loja, and El Oro Provinces. Its elevational range is 1682–3018 m in an area of extent of occurrence of about 1621.2 km 2.
This nocturnal, semiarboreal species inhabits mostly disturbed areas and a few forests in the Evergreen Montane Shrub in the south of the Ecuadorian Andes, the Evergreen Montane Forest of the Cordillera Occidental of the Andes, and the Evergreen Montane Forest from the south of the Cordillera Oriental of the Andes (Ministerio de Ambiente del Ecuador 2012), where the average annual rainfall is 566–1066 mm and the average annual temperature is 12.2–18.8 °C (Fick & Hijmans 2017). At Oña, Azuay Province, several Gastrotheca lojana were calling from agave leafs 50–130 cm above the ground and among grasses near the ground (LAC field notes, 8 August 2003). A male from San Fernando, Azuay Province, was calling from approximately 100 cm above the ground on a large stone. It was close to another adult and a juvenile of the same species; the stone was 7 m from a pond. Another individual was among totora reeds (Schoenoplectus californicus) approximately 8 cm above a small stream. A brooding female is depicted in Figure 10L. Throughout most of its distribution, G. lojana is syntopic with G. pseustes; at San Fernando, it is syntopic with G. pseustes and G. litonedis, and in the southern part of its range it is also syntopic with G. elicioi.
Conservation status. We suggest that Gastrotheca lojana should be considered as Endangered according to criteria B1ab(iii,v) of the IUCN Red List. We suggest this category because its small area of occurrence (1621 km 2) is fragmented, and its habitats are in heavily human populated areas. Several search efforts (in 2010–2013, 2016, 2017) at Loja and surroundings revealed no specimens. Environs of Laguna de Busa and Oña have been deforested and modified for human activities (8D, E). The Busa lake is populated by introduced fish (trout and carp) and its surroundings by introduced eucalyptus and conifers. Additional threats at Laguna de Busa are unregulated tourism activities. None of its populations is included in the National System of Protected Areas (SNAP).
Comments. Duellman (1974) referred some specimens from 10 km W Loja, (KU 142603–8, 148549–51), Loja Province, to Gastrotheca lojana. These are identified here as G. elicioi (see below). Also we identify as G. lojana specimens from Girón, Azuay Province (KU 138401–138403), and Saraguro, Loja Province (KU 138404– 138409) that Duellman (1974) referred to G. monticola. In Duellman & Hillis (1987), specimens of G. lojana were also included in the description of G. monticola. Loaiza-S (2012) provided a summary account of G. monticola, in which G. lojana was included. Duellman et al. (2014) and (Duellman 2015) recognized G. lojana as genetically distinct from G. monticola. Duellman (2015) provided an account of G. lojana. A specimen from Oña, Azuay Province (QCAZ 2692) depicted as G. litonedis in Duellman (2015:239, Fig. 11.12 B) belongs to G. lojana.
description: Published as part of Carvajal-Endara, Sofía, Coloma, Luis A., Morales-Mite, Manuel A., Guayasamin, Juan M., Székely, Paul & Duellman, William E., 2019, Phylogenetic systematics, ecology, and conservation of marsupial frogs (Anura: Hemiphractidae) from the Andes of southern Ecuador, with descriptions of four new biphasic species, pp. 1-102 in Zootaxa 4562 (1) on pages 71-79, DOI: 10.11646/zootaxa.4562.1.1, http://zenodo.org/record/2627982