Natural History and Intraspecific Variation of the Ecuadorian Blue Glassfrog Cochranella Mache GUAYASAMIN & BONACCORSO, 2004 (Anura: Centrolenidae)
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HERPETOZOA 21 (1/2): 57 - 66 57 Wien, 30. Juni 2008 Natural history and intraspecific variation of the Ecuadorian Blue Glassfrog Cochranella mache GUAYASAMIN & BONACCORSO, 2004 (Anura: Centrolenidae) Biologie und innerartliche Variabilität des Ekuadorianischen Blauen Glasfrosches Cochranella mache GUAYASAMIN & BONACCORSO, 2004 (Anura: Centrolenidae) D. F. CISNEROS-HEREDIA & J. DELIA & M. H. YÁNEZ-MUÑOZ & H. M. ORTEGA-ANDRADE KURZFASSUNG Dia Autoren präsentieren neue Erkenntnisse zur Biologie und innerartlichen Variabilität von Cochranella mache GUAYASAMIN & BONACCORSO, 2004. Die nachtaktive Art kommt entlang kleiner Bäche in Primär- und alten Sekundärwäldern der Cordillera Mache-Chindul vor, und ist ein Endemit der saisonal immergrünen Wälder West- Ecuadors. Die intraspezifische und geschlechtsdimorphe Variabilität zahlreicher Merkmale einschließlich der Textur der Rückenhaut, der Ausprägung von Hautfalten, Kloakalarmatur und Supratympanalfalte sowie der Aus- dehnung der Iridophoren auf dem Leberperitoneum, wird dargestellt. Es wird über einen tiefgreifenden, für Centrolenidae einzigartigen Farbwechsel berichtet, bei dem das Farbmuster des Rückens von bläulich-grün mit einem blaßgelben Fleck auf dem Kopf und zahlreichen gelben Tupfen zu einer lavendelfarben-hellblauen Grund- farbe mit hellgelbem Kopffleck und zahlreichen orangefarbenen Punkten wechselt. Um künftigen Untersuchern hilfreiche Informationen zur Charakterisierung der Art an die Hand zu geben, werden die neuen, für Co. mache dia- gnostischen Merkmale übersichtlich aufgezählt. ABSTRACT We present new information on the natural history and intraspecific variation of Cochranella mache GUAYASAMIN & BONACCORSO, 2004. It is a nocturnal species associated with small streams across primary and old- secondary forests in the Cordillera Mache-Chindul, endemic to the Seasonal Evergreen forests of the West Ecuadorian Region. Intraspecific and sex-related variation is noted in several characters, including dorsal skin tex- ture, expression of the dermal folds, cloacal ornamentation, expression of the supratympanic fold, and extent of iri- dophores on the hepatic peritoneum. Dramatic chromatic changes in life are reported, showing a unique pattern in Centrolenidae, its dorsal pattern changes from bluish-green with a dull yellow patch on the head and abundant yel- low spots, to a lavender/light blue dorsum with a bright yellow patch on the head and abundant orange spots. In order to provide future researchers with useful information to characterize the species, we present a numbered diag- nosis for Co. mache that includes all new information. KEY WORDS Amphibia: Anura: Centrolenidae; Cochranella mache; morphological variation, morphology, coloration; sexual dimorphism, systematics, biology, natural history; Ecuador INTRODUCTION The glassfrog Cochranella mache HEREDIA & MCDIARMID 2007). Cochranel- GUAYASAMIN & BONACCORSO, 2004 was de- la mache is considered a threatened species scribed from three male specimens collect- and remains poorly understood (GUAYA- ed at one locality on the eastern slope of SAMIN 2007). Recent surveys carried out in Cordillera Mache–Chindul (= Montañas de the province of Esmeraldas, Ecuador, have Mache), a rather isolated mountain range in yielded new information on Co. mache in- the northern portion of Cordillera de la cluding data on aspects of its natural history Costa in western, coastal Ecuador (GUAYA- and intraspecific variation, which we pres- SAMIN & BONACCORSO 2004; CISNEROS- ent herein. 58 D. F. CISNEROS-HEREDIA & J. DELIA & M. H. YÁNEZ-MUÑOZ & H. M. ORTEGA-ANDRADE MATERIALS AND METHODS Characters and terminology follow the third finger (3DW), tibia length (TL), definitions and proposals by CISNEROS- and foot length (FL). The following abbre- HEREDIA & MCDIARMID (2007). Termino- viations are used along the text: Ce. = Cent- logy for the webbing formula follows the rolene, Co. = Cochranella, H. = Hyalino- method of SAVAGE & HEYER (1967) as mod- batrachium, and N. = Nymphargus. The ified by SAVAGE & HEYER (1997) and latter three Glassfrog species from southeastern summarized by GUAYASAMIN et al. (2006) Brazil and northern Argentina were exclud- and CISNEROS-HEREDIA & MCDIARMID ed from Hyalinobatrachium by CISNEROS- (2007). Sex was determined by direct exam- HEREDIA & MCDIARMID (2007) and left as ination of the gonads and by noting the pres- incertae sedis. They are listed here as ence of secondary sexual characters (i.e., “Hyalinobatrachium” eurygnathum (LUTZ, vocal slits, nuptial pads). Internal anatomy 1925), “Hyalinobatrachium” parvulum was examined by dissection of recently (BOULENGER, 1895), and “Hyalinobatrachi- euthanized specimens. The following um” uranoscopum (MÜLLER, 1924) to de- measurements and their abbreviations are note that they are not formally attached to cited in the text; all were taken with elec- any genus within Centrolenidae. Institu- tronic digital callipers (0.05 mm accuracy, tional abbreviations used are as follows: rounded to the nearest 0.1 mm) as described DHMECN — División de Herpetología, by CISNEROS-HEREDIA & MCDIARMID Sección Vertebrados, Museo Ecuatoriano de (2006, 2007): Snout-vent length (SVL), Ciencias Naturales, Quito; DFCH-USFQ — head width (HW), head length (HL), hori- Universidad San Francisco de Quito, Quito. zontal eye diameter (ED), inter-orbital dis- Classification of vegetation formations in tance (IOD), eye-nostril distance (EN), Ecuador follows SIERRA (1999) as modified internarial distance (IN), width of disc of by CISNEROS-HEREDIA (2006; 2007). RESULTS Expeditions to three localities in the Canandé is the first known female of the Province of Esmeraldas, Ecuador, found species. It had unpigmented eggs and was one specimen of Cochranella mache each: collected at the end of the local dry season. DHMECN 2611, an adult male collected at No reproductive activity (calls, territorial night while perched on a herbaceous plant males, or females with eggs) was observed about 1.6 m above a small stream at Monte for any other centrolenid at the site at that Saino, Punta Galeras region (00º42’N, time. When found, the female was using 80º01’W, 100 m elevation), on 21 October its hind leg to kick an approaching cock- 2004; DFCH-USFQ LQ23, an adult male roach. Cochranella mache was sympatric found at night perched on a low broadleaf with Centrolene prosoblepon (BOETTGER, shrub about 1 m above a small stream in 1892), Ce. ilex (SAVAGE, 1967), Co. albo- old secondary forest about 3 km NW from maculata (TAYLOR, 1949), and an unde- the town of Quininde (00º21’N, 79º29’W, scribed species of Cochranella at Canandé; about 150 m elevation), on 23 March 2005; with Ce. prosoblepon, Co. pulverata (PE- and, DHMECN 3560, an adult female TERS, 1873), and Hyalinobatrachium fleisch- encountered at night perched on broadleaf manni (BOETTGER, 1893) at Monte Saino; herbaceous vegetation (0.87 m above for- and with Ce. prosoblepon, Ce. litorale est floor) about 2.30 m from the edge of a RUIZ-CARRANZA & LYNCH, 1996, Co. albo- small stream (depth 0.11 m; width 0.89 m) maculata, and H. fleischmanni at the sur- in old secondary forest at the Reserva roundings of Quinindé. At its type-locali- Biológica Canandé (00°26’N, 79°08’W; c. ty, Bilsa, Co. mache was sympatric with 270 m elevation), on 24 June 2005. The Ce. prosoblepon, Co. albomaculata, and specimen (DHMECN 3560) collected at H. fleischmanni. Natural history and intraspecific variation of Cochranella mache 59 Table 1: Variation of measurements (in mm) of one female and two male specimens of Cochranella mache GUAYASAMIN & BONACCORSO, 2004. For abbreviations see ‘Materials and Methods’. Tab. 1: Morphologische Meßwerte (in mm) bei einem Weibchen und zwei Männchen von Cochranella mache GUAYASAMIN & BONACCORSO, 2004. Abkürzungen siehe ‘Materials and Methods’. Parameter DHMECN 3560 DHMECN 2611 DFCH-USFQ LQ23 Female / Weibchen Male / Männchen Male / Männchen SVL 28.0 22.0 23.5 HW 8.4 7.5 7.6 HL 8.4 7.6 7.7 ED 2.7 2.7 2.6 IOD 3.8 3.4 3.5 EN 2.7 2.1 2.3 IN 2.6 1.7 1.6 3DW 0.9 0.9 0.9 TL 16.3 13.8 14.1 FL 13.6 11.2 11.6 The original description of Cochranel- tubercles, but sometimes becoming so la mache (based solely on males, GUAYA- inconspicuous that just the tubercles are SAMIN & BONACCORSO 2004) is adequate, apparent (dermal folds present, conspicu- thus we will not repeat general details about ous, with large white tubercles on ventrolat- the species’ morphology. Instead, we pro- eral edges of finger IV, forearms, elbows, vide information on characteristics of the toe V, tarsi, and heels); (3) cloacal opening first female specimen known, additional bordered laterally by low, fleshy, tuberculat- variation observed, and morphometric and ed horizontal flaps but not n-shaped; flat, chromatic accounts. The adult female is large, fleshy cloacal enameled tubercles similar to males in most characters of their located posterior to cloacal slit and towards general morphology. The main differences the vent (cloacal opening bordered laterally include (male characters in parentheses): (1) by fleshy tuberculated n-shaped fold; cloa- dorsal skin shagreen and with warts but cal tubercles small, fleshy, located immedi- without spicules (dorsal skin shagreen, usu- ately posterior to cloacal slit); and (4) white ally with numerous minute spicules and coloration restricted to the points of finger I warts with protruding spicules); (2) ventro- and II, points of toes I and II, but also on the lateral edges of fingers IV, forearms, margins of webbing between fingers and elbows, toe V, tarsi, and heels